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VOL. 71 PART 1 

JULY 25, 1947 






Fifteen Shillings 

Registered at the General Post Office, Adelaide, 
for transmission by post as a periodical 

VOL. 71 — 1947 





Registered at the General Post Office, Adelaide, 
for transmission by post as a periodical 


Womersley, H., and Kohls, G. M. ; New Genera and Species of Trombiculidae from 

the Pacific Islands . . . . . . . . . . . . . . 3 

Johnston, T. H,, and Edmonds, S. J.: Australian Acanthocephala No. 5 .. .. 13 

Black, J, M. : Additions to the Flora of South Australia, No. 44 20 

Johnston, T. PL, and Mawson, P. M.: Some Nematodes from Australian Lizards r . 22 

Turner, A. J.: A Revision of the Australian Phycitidae, Part I 28 

Miles, K. R. : Pre-Cambrian Granites and Granitisation, xvAh special reference to 

Western Australia and South Australia . . . . . . . . . . 54 

Prescott, J. A., and Lane-Poole, C. E. : The Climatology of the introduction of Pines 

of the Mediterranean Environment to Australia . . . . . . . . . . . . 67 

Crocker, R. L., and Wood, j. G. : Some Historical Influences on the Development of 
the South Australian Vegetation Communities and their bearing on Concepts and 
Classification in Ecology . . . . . . . . . . . . . . 91 

Burbidge, Nancy T. : Key to the South Australian Species of Eucalyptus L'Herit .. 137 

Stephens. C. G. : Functional Synthesis in Pedogenesis .. .. ,. .. .. 168 

Finlayson, H. H. : On the Weights of some Australian Mammals 182 

Wilson, Allan F. ; The Charnockitic and Associated Rocks of North-Western South 

Australia. Part I, The Musgrave Ranges — an Introductory Account , . . . . . 195 

St'Rroc, Reg. C. : Early Cambrian ( ? ) Jellyfishes from the Flinders Ranges, South 

Australia 212 

Eva\s, J. W. : Some New Eurymelids from Australia and New Guinea ( Homoptera, 

Jassoirica) . . . . . . . . . . . . . . . . . . . . . . 225 

Wo"mkrslf.y, IT. B. S. : The Marine Algae of Kangaroo Island. I. A General Account 

of the Algal Ecology 228 

Kk'klinx, L. C. P.: Pythium debaryanum and Related Species in South Australia .. 253 

Mawson, D. : The Adelaide Series as Developed along the Western Margin of the 

Flinders Ranges . , . . . . . . . . . . . . . . . . - . . . 259 

La\'gford-Smith, T. : The Geology of the Jamestown District, South Australia .. .. 281 

Sl'RIGG, Reg. C. : Submarine Canyons of the New Guinea and South Australian Coasts . . 296 

AVomersley, H. E. S. : The Marine Algae of Kangaroo Island, f. A General Account 

Western Australia 311 

Ohhei,, I. L. : Notes on the Genera Lychnothamnus and Lamprothamnium (Characeae) 318 

Johnston, T. Harvey, and Beckwitii, Anne C. : Larval Trematodes from Australian 

Freshwater Molluscs. Part XII 324 




Cecil Thomas Madigan, M.A., B.E., D.Sc, F.G.S., a leading Australian 
geologist and geographer, a member of this Society for the past 25 years, and a 
Past President, died at the early age of 57 years on 14 January 1947. 

Curtailment of his span of life was doubtless the result of heart strain result- 
ing from exceptional exertions in the field of sport, exploration and war. 

His earlier education was received at the Adelaide High School, Prince 
Alfred College and the University of Adelaide, where he graduated in Mining 
Engineering in 1910, As a Rhodes Scholar from South Australia, he then pro- 
ceeded to Magdalen College, Oxford, to continue with geological studies, which 
were, however, to be postponed, for late in 1911 he joined the Scientific Staff of 
the Australasian Antarctic Expedition of 1911-14. 

Dr. Madigan's record during more than two years in Antarctica was out- 
standing. It includes leadership of a sledging party which reconnoitred the ice 
plateau in winter time under record adverse climatic conditions. In the summer 
of 1912-13 he successfully led a sledge journey, charting a great length of new 
coast line of the territory thereafter known as King George Land. On return 
to winter quarters, he was selected to be in charge of all land operations pending 
the return of the Expedition Leader, who at that time had failed to return from 
a journey across the plateau ice. 

His graphic account of the summer sledge journey forms a section of the 
popular story of the Expedition, published as "The Home of the Blizzard." He 
was chiefly responsible for the meteorological record of the Cape Denison Station, 
which is published as an important section of the Expedition's Scientific Reports 
issued from the Government Printing Office, Sydney. 

On the advent of war when again in England in 1914, he joined the Royal 
Engineers and served throughout the whole period with rank of Captain. Later, 
having finally graduated at Oxford, he was appointed Assistant Government 
Geologist in the Soudan. This post he relinquished in 1922 to join the staff of 
the University of Adelaide. 

With the outbreak of the Second World War, Dr. Madigan was appointed 
chief instructor in the School of Military Field Engineering (Liverpool, N.S.W.), 
attaining the rank of Lieut-Colonel. 

During his long term as lecturer in Geology, he found time to execute a 
number of important geological investigations dealing with problems relating to 
South Australia and to Central Australia. It is in the latter field that his work 
has received most recognition. His explorations in the MacDonnell Ranges and 
neighbourhood, and in the Simpson Desert, have established him as the leading 
authority on the Geography and Geology of Central Australia. For this work he 
was the recipient of wide recognition in scientific circles both in Australia and 
abroad. He is also well remembered for his zeal in furthering the good work 
of the Legacy Club and the University Graduates' Association. 

His published scientific works additional to those dealing with his Antarctic 
activities mentioned above are the following: — 

"A Description of some Old Towers in the Red Sea Province, North of Port 

Soudan." Soudan Notes and Records, 5 (1922), 78-82. 
"The Geology of the Fleurieu Peninsula, Part I — The Coast from Sellick's Hill 

to Victor Harbour." Proc Roy. Soc. S. Aust, 49 (1925), 198-212. 
"Organic Remains from below the Archaeocyathinae Limestones at Myponga 

Jetty, South Australia." Trans. Roy. Soc. S. Aust., 50 (1926), 32-33. 
"The Geology of the Willunga Scarp." Proc. Roy. Soc. S. Aust,, 51 (1927), 

A.N.Z.A.A.S. Reports of the Glacial Research Committee (with D. Mawson), 19 

(1928), 97-99. 
"Preliminary Notes on New Evidence as to the Age of Formations on the North 

Coast of Kangaroo Island." Trans. Roy. Soc. S. Aust., 52 (1928), 

"An Aerial Reconnaissance into the South-Eastern Portion of Central Australia." 

Proc. Roy. Geog. Soc. (S. Aust. Branch), 30 (1929), 83-108. 
"Lake Eyre, South Australia." Geog. Jour., 76 (1930), 215-240. 
"Pre-Ordovician Rocks of the MacDonnell Ranges, Central Australia" (with 

D. Mawson). Qld. Jour. Geog. Soc, 86 (1930), 415-428. 

"Geology of the Western MacDonnell Ranges, Central Australia." Qld. Jour. 

Geog. Soc, 88 (1932), 672-711. 
"The Geology of the Eastern MacDonnell Ranges, Central Australia." Trans. 

Roy. Soc. S. Aust., 56 (1932), 71-117. 
"The Physiography of the Western MacDonnell Ranges, Central Australia." 

Geog. Jour., 78 (1932), 417-433. 
A.N.Z.A.A.S. — Reports of the Glacial Phenomena Committee, 21 (1932), 464. 
"The Geology of the MacDonnell Ranges and Neighbourhood, Central Australia." 

Reports A.N.Z.A.A.S., 21 (1933), 75-86. 
"The Australian Sand-ridge Deserts." Geog. Rev. (1936), 26, (2), 205-227. 
"Central Australia." Oxford University Press, 1936 and (2nd ed.) 1946. 
S. Aust. Royal Society Centenary Address : "The Past, Present and Future of 

the Society, and its Relation to the Welfare and Progress of the State." 

Proc Roy. Soc. S. Aust., 60 (1936), I-XV 

"Additions to the Geology of Central Australia." A.N.Z.A.A.S. Reports, 23 

(1937), 89-92. 
"A Review of the Arid Regions of Australia and their Economic Potentialities." 

A.N.Z.A.A.S. Reports— Presidential Address, Section P, 23 (1937), 375 

"The Boxhole Crater and the Huckitta Meteorite." Proc. Roy. Soc S. Aust., 61 

(1937), 187-190. 
"The Huckitta Meteorite, Central Australia" (with A. R. Alderman). Min. Mag., 

(1939), 25, (165), 353-371. 

"The Boxhole Meteoritic Iron, Central Australia" (with A. R. Alderman). Min. 
Mag., (1940), 25, (168), 481-486 

"Simpson Desert Expedition, 1939: Scientific Reports — Introduction, Narrative, 
Physiography and Meteorology." Proc. Roy. Soc. S. Aust., 69 (1945), 

"Simpson Desert Expedition, 1939: Scientific Reports — No. 6 — Geology: 
The Sand Formations." Proc Roy. Soc. S. Aust., 70 (1946), 45-63. 

Adelaide, 14 July 1947 D. M. 



ByH. Womersleyand GM.Kohls 


Description of Larvae- Shape ovate. Length (engorged) to 450, width to 300. Scutum pentagonal, as 
figured, with the anterior margin lightly sinuous, posterior angle rounded. Sensillae ciliated on 
distal half, with the sensillae bases nearer to each other than to PL and placed slightly anterior of 
line of PL. Eyes 2+2, on distinct ocular shields, the posterior eyes the smaller. Chelicerae not 
serrated on inner (dorsal) edge, with the usual small apical tricuspid cap forming the apical tooth, a 
small subapical dorsal tooth, and a small subapical ventral tooth. Galeal setae nude. Palpi stout, 
tibial claw bifurcate; seta on palpal femur 2-3-branched; on genu nude, on tibia only the ventral seta 
branched; tarsi short, with basal and subapical rods, and five or six ciliated setae, one of which is 
much stronger than the rest and over-reaches the tip of the tibial claw by the claw's length. 



By H. \ (i: ' and G. M, Konu; ( -> 
[Read 10 October 1946] 

Eutrornbicula gymnodactyla if. sp. 

Fig, 1 A-F 

Description of Lo?V$& — ShajyS ovate. Length (engorged) to 450 /i, width 
to 300 /a, Scutum pentagonal, as figured, with the anterior margin lightly sinuous, 
posterior angle rounded. Scnsillae ciliated on distal half, with the sensillae bases 
nearer to each other than to PL and placed slightly anterior of line of PL Eves 

Fitr- 1 Eutroviblcufa (Asta-nscus) fjxmnoiiaclyfo v.. <\>. A, dorsal scutum 

(x500); H, palp in dorsal view (x 860) : C, tt;i of e.hplkera (\-fi60); D, galeal 

seta; F, tarsus &ad metatarsus tit (x 450) ; F, dorsal seta O860). 

2~\-2, on distinct ocular slrelds. the posterior eyes the smaller. Chelicerae not 
serrated on inner (dorsal) edge, with the usual small apictd tricuspid cap form- 
ing \\\e apiea; tooth, a small suhapical dorsal tooth, and a small subapieal vcnrral 
tooth. Gajeal setae mice. Palpi stout, tibial claw bifurcate . seta on palpal femur 
2-3-branched; on genu nude, on tibia only the ventral seta branched; tarsi short, 
with basal and subapical rods, and five or six ciliated vSetae, one of which is much 
stronger than the rest and over-readies the tip of the tibial claw by the claw's 

Dorsal setae 24 in number, to 3G /t long, ciliated auc arranged 
VcutralJy with a pair of branched setae on guathosoma, one;, pa each coxa, a pair 
between coxae T and betwee.n coxae ilT, thereafter uncertain as all specimens 

C 1 ) Entomologist, South Australian Museum, 

C 2 } Major S'-i.C-, Upucd States oi America Typhus Coiimiisinoti, now Entomologist, 
"[T.S- Public TTca'tb Service, "Rocky Moif-nnm Laboratory, Hamilton, Montana. 

Traus. Key. Soe. S. Aiist, 7J (1). 25 July i 04? 

were heavily im^ied with sporozoa but approximately to 20 n in length. Legs.: 
I 210 {x long. 11 195//,. Ill 210/;.; tarsi 1 and II with the usual dttrsfj sensory 
rods; tarsi III without any long nude beta. 

The Standard Data (s&e Wottievsley and li'easlip 1945) in microns for thc 
tvpe and three paratypes. 

' L Standard 'I 'Tipoff Ifcnl ONscrvfcd CV,ei>\ 01 

Mvv.u Dcvi;itic:i JOn^i; * l&a&t Variation 

AW - - 45-75^0-75 1-50^0-50 41-2." 50-25 45-0-48-Q 3-2 

PW . - < . U - No variation p&aCaed 

SB ... kho rt0-87 1-73+0-61 Il-sd -gt>7 15-0-13-0 10*5 

ASB lo'O No variaiion records -J 

PSB . ZJ«S5±SW5 l-5(.i±0*53 IJMi 23-75 21 -0— 24-0 6-4 

SD .... 4i-25dr0-75 l-SOrtO-S^ 8(5 v3- 4W3 39 -0-42-0 ivi) 

A-P L'-5 drf! * H7 l-7Jrb'>6L 14-5 2-1-7 1U*Q— 21*0 8-8 

A.AI ... 24-0 No variation recorded 

AL ... 2.5-5 ±0-S7 l-73±0-61 20 -3 -50-7 2!-0 27-0 G-8 

PL 21-0 No variation recorded 

$m*'~ 47-0 ±1-0 2-73—0-71 41-H-52-2 4"-0-4a-0 3-7 

/^ ;r . Th ; 5 species L describe! from tour specimens from Cyinnodartyhts 

lQuisk?dv)isis from Sansipor. Dutch New Guinea (G. §1 K_. Sansapor, No. 3, 
26 Aug. 1944). 

Remarks— In the pentagonal srtuum th : s species would fall into S.ig Triors 
subgenus Peutarjoneih 1935, but the bifurcate p ia ipal daw puts it jnto^ Ewdng^ 
Euirov.ihlcvJa 1938, and the number at dorsal s/etae into Ascarisrus 3£vvmg 1943. 
We concur, however, v.dlli Miehenc r 1946 (Anna is. LnL Soe. America, 29, (1), 
101 118, iu regarding Asrarisc:;s as a synonym of Enirautbiniia. 

The type and one puvalyne in the South Australian Museum, and two para- 
topes hi th:e U.S. National Museum. 

Ceuus Novofcrcmbicula nov. 
L//frae— As iu TriVKhiruIa Berk si, with the dorsal scutum furnished with 
paired filamentous scnsnku} and the usual five U\*£ ( niiP A -'^ twoAL, and two 
PLL b'ut produced posteriorly and tongue-like, so as to include, in addition to 
the above, some of the median setae of some of the dorsal rows as in the gtSflU 

Genotype Ivoisoinirtulnatfa ozv!cns;s n. sp. 

Navotrcmbicula owiensrs jji &yr. 
Fig; 2A-K 
Ucnnl'lhn of f^rrt^— Shape subroiund. Size snlall ; length (unfed) 255 /*, 
Width ioO/*. Dor^l seutnni large, tongue-shaped and produced posteriorly to 
;ake hi the two median setae of IlYe third dorsal row, in addition to the AM, AL 
and PL setae; scnsillae lone; and filamentous with i-rialions on the distal half. 
bases wide apart ami about midway between Eiws of AL and PL. Eyes 2 + 2, 
on d'-stinet ocular shields placed close to the lateral semal margin ; posterior eyes, 
ike smaller, Cheiicera with only the raiurJ apical tricuspid cap, Int. on the inner 
taagni W&ft the cap with a rather prominent suhapka! angle. Grikal setae long 
nv.a niulo. Pnlpi rather lender, with rather fcttg skndcr and bifurcate tibial 
claw, Ihe dorsal" prong of which is email and indistinct; palpal femur stid genu 
with' a iomr slender,** shortly cdiated seta; all three setae on tibia long and 
apparently nude; tarsi moderately loflg and couica', with a fting sithba*al sensory 
rod -ud iive or six lun- ciliated scUtc 

Dorsal setae rather thick, slightly tapering, shortly ciliated and of two sizes, 
arranged 2.4.4 [2]. 4.2.2; the second row with the medial pair just off the scutum 
and to 57 n long; the inner members of the third row are on the scutum and are 


1 : ■ it! A \ i 

"^ Ms 

■7 \ £*? 



/i 4 > 

/ vp ir 1/ \ 

A I H*w^ V W \ \ 


v * \ *- •*■ i 


1 r-j 



\ 1 Mi , 

./ \ 

\ t 





J ■*-- — ■- ^ 

f A 

f $ 

/ \ 




9 ' 


/ 1 



>■•■■■ * B _ ^ - : ^t:_ _ 

Fig 2 Kovotrombicufa ozvlcr.sis n. K*. J3ld n. ?p. A, ccrsal view; B, ventral 
view; C, dorsal scutum (x 500) ; D, palp in dorsal view (x 860) ; E, tip of 
chelicera (x 860) ; F, galeal seta; G, dorsal seta (x 860) ; H, tarsus and meta- 
tarsus III (x450). 

short to 24 fi. Ventrally with a pair of branched or ciliated setae on the gnatho- 
soma. one on each coxa, a pair between coxae I and between coxae IIT and there- 
after to 24 fi long. All three pairs ol coxae touching. Legs: I 255 /a 
long. 11 240 /a 111 275 fx; tarsi T and 11 with the usual sensory rod-like sela, 
III without any long nude seta. 

The Standard Data in microns for type and two paratypes are : 


Th* [rnrgjiiifti 


Cocft. of 






A W , . 



83-1 9sV5 

87-0- 91-0 


nv . 

95 -3 ±1-33 

2-31 +0-94 

88-0— 102-2 

94 -0- 9&-0 


ST3 ... 

61 -ft+rl -00 


55 •#— '.6-2 

6(1-0— 63-0 


ASB ... 


1- /o±0 -71 

Jfl-R- 21-2 

15-0— 18-0 



112-0 No 

vanation recorded 

SD . 




127-0— 130-0 


A l> .. 



19-1— 22-3 

20-0- 21-0 



. 40-3-0-88 


35«7— 44*7 

39-0- 42-0 



JQ-fl No 

variation record. 


PL ... 

4^0 Xo 

variation record; 


Sen?. . 

. 67-0:fcl-0 


6t-g— 72 '2 

6G-C— 69-0 


Loc— Six specimens collected from the soil at the base or a large tree oil 
Owi Island, North Coast of Dutch Xew Guinea. 15 Aug;. 1944 (G. M. K., 
Owl 22 C). 

h'ewnrks — But for the prolongation of the dorsal .scutum to include some of 
the medial dorsal setae, this interesting species would fall into Tromhicida s.l. 
The form of the scutum, however, is parallel to that found in the genus Gahrliepla 
of the group of genera with clavatc sensillae and justifies the erection of a new 

The lype and two paratypes in the. Soulh Australian Museum, three para- 
types in the collection of the Rocky Mountain Laboratory, 

Schongastia philipi n. sp. 

Fig. 3 A-G 

Description cf Larvae — Shape an e'ongate ovah Length (unfed) 236 fx, 
width 182 fx, Scutum more or less hexagonal, with the posterior margin 3-sided 
and deep behind line of PL with a slight concavity medially; AM seta very much 
shorter than AL and PL. AL the longest; sensillae more capitate than clavate 
with their bases in line with PL. Lyes 2 -j- 2. apparently not on ocular shields, 
the posterior eyes the smaller. Chelicerae not unusually slender, with apical 
tricuspid cap and on the inner (dorsal) margin with three to four strong and 
some smaller teeth. Galea! setae nude. Palpi si out,, with trifurcate tibial claw; 
seta on palpal femur ci'iated or branched; on genu mule; on tibia dorsal and 
lateral nude, ventral branched; tarsus short, with basal and subapical sensory 
rods and several ciliated .setae. 

Dorsal setae 36-40 p> long, ciliated. 28 in number and arranged 
Ventrally w?th paired branched seLae on gnathosoma, a ciliated seta on each coxa, 
a pair between coxae 1 and between coxae III, and thereafter to 25 /i 
Ujng, Legs: I 256 /v, long, Tl 228 /a, ITT 256//.; tarsi I and IT with dorsal sensory 
rod* ITT with a long nude seta arising; sttbbasally and dorsally. 


The Standard Data lor type and four paratypcs arc: 


Mean Deviation Jiange 

AW .... 4S-6±1-12 2-51±0-80 41*1 56-1 

P W -■■ &* feO '48 - 89-±0 -28 66-9- ~?2 ■ 3 

SB ... 18-4:h0-40 0-89±0-28 15-7—21-1 

ASB .... 24-0 No variation recorded 

PSB .... 21-0 No variation recorded 

SD .... 43-0 No variation recorded 

A-P ... 23«f>±0-40 0-89±0-28 20-9— 26-3 

AM .... 22'2r±0-73 1-64^0-52 17-3— 27-1 

AL ... 59-2±0-49 l-04±0-34 55-9— 62-5 

PL - - 42-6±:M2 2-5l±0-79 35-1-50-1 

Sens oO'O with head 15/18 

Ol: set red 




45-0 51-0 


OS -0-70-0 












\ i » ; 

Pig. 3 Schongast hi philipl n. sp. A, dorsal view; B, ventral view; C, dorsal 

scutum <x50U); T) t palp m dorsal view (xS60); E, tip oi chelicera (x86U); 

P, galeal seta; G, tarsus and metatarsus III (x450). 


Loc,— r l'ypv, and nine paratypes from a lizard LeiolcjAsnia uibrrilsn from 
Goodenough Island, New Guinea. 17 Jan. 1944 (G. M. K., No. 4S6). 

ticnutrks — The dentition of the. cheh'ccrac in this species is somewhat different 
from the many small teeth found in other .species of SchotufardiO; but this is not 
fetiftidern to warrant generic separation. 

The type and lour paratypes in the South Australian Museum, and one para- 
tope to be deposited in Ui$ British Museum; one puratype Lo be deposited in the 
C.S, .National Museum and in the Rocky Mouuiudn Labcr^iory. 

Genus Ocnoschcngattia nov. 
Allied to Schbnrjastl:: hut the ehcliecrac are short, cm ve 1 and vlnmpy with 
the apex divided iuio two terminal b'um and thick teeth, with a pjur (.-xanetimes 
three) of rather smaller teeth subapieaifv. Palpal claw short and stout, tri- 
furcate. Median tarsal claw (empoduun) longer than the laterals but equally 
thick. Fensillac clavate or capitate. 

Qenoschongastia cana n. sp. 
Fig. 4A-G 

lh\~criptioji of Larvae— Shape au elongate oval. Length (unfed) 252 /*, 
width 162 f.L. Dorsal scutum roughly hexagonal with iudlstinet and fine pitting: 
with the usual five normal cibaled seta^ t of which AL are [he tottgeM and AM 
the shortest, AM with short hranches, AL and VL with long outstanding 
branches; sensiliae broadly clavatc, the head indistinctly and very shortly sctulose, 
sensillae bases about in line Tvifcft PL; anterior scutrJ margin convex, posterior 
laterally angular, Ryes 2 -(- 2. large, on well-developed ocular shields and closely 
adjacent to scutum, posterior eyeg the smaPer. Chclkerae of peculiar form, short, 
stout and curved, apex truncate and divided into two siroug. broad and blunt 
teeth, and two fscmetiuies three) smaller teeth placed subapleally on the inner 
edgc. Galeal setae mtclc. Palpi stout, tibial claw short, stout and trilurcate; 
femur with a l&ug strong &?i# with long branches; £enu with a nude seta; tibia 
with the dorsal and hueral setae nude, ventral branched; tarsi short with basal 
and subapieai sensory rods and three or four ciliated or branched setae. Dorsal 
setae strong with strong ciliations, 36 in number and arranged, to 
40-50/4 Ions'- Ventrally with the usual pair of branched setae on guathosoma, a 
single sela on each coxa, a pair between coxae I and between coxae J It, and there- 
after, more slender and with longer citations than the dorsal setae, to 
30 /.i Iou^. Legs: I 324// long. II 252 ji. Ill 3d0/.i; tarsi I and II with the usual 
dorsal rod-like se'.ae, tarsi ill with a long outstanding nude seta* ; median claw 
(empodium) longer than but as ihiek as the laterals. 

The Standard Data in microns derived from 22 specimens are: 




fneff- of 







, 62*85^0-56 


54-9 -70 -H 




. 7S-7 £0*71 


<V>7 S3 -7 

75-0- 87-0 

A -2 





:ir>G— 3'ho 


ASP ... 

. 2$-V ±9i$i 


t$4 ~33>o 

27-0 :2-0 


Z\} *j 

a^-«a -±0'.3o 


KWi — 2S-Q 




. 52-8 dt0*45 

2-M + Q-32 

-!6-5 - :o-i 



A-;-* ... 

29-2 ±0-32 


|4>7 ~^>7 

27-0— 3 &* 


A\L ... 

. 32-45;h0-!6 


2fr0 -:)$•<) 

27 -0 -35-0 


A', , 

74-3 ±OoO 


c/-3 -Jrjg 



?L .. 

. 59-5 ±0-o9 


?|M) —65-0 

55-0 ifS-iJ 


Sern. .. 

s>o-0' willi 

head 20724 

* Unfortunately omitted from fig. 4 G. 

Loc. — The type and 72 parritypes from the mound oi a brush Lurkey, Dobo- 
dura, New Guinea, IS May l&f<f £& if. K, No. 325). 

The type and 22 paratype deposited in the Soutli Australian Museum, 35 
paratypes in the collection of ihc Reeky Mountain Laboratory, five paratypes m 
the U.S. National Museum and live in the British Museum. 

J-h'VMirks— This genus and species in the peculiar and characteristic cheliccrae 
and the palpal ciaw is very distinct from Schovgasiht, Ncoschongasiia and 


t r-V%& 

ffl III 1/ 1/ ! 

\ ft 1,4$ ' I Mw / / 

■\ i . L mm .y // a> // if ■■"/ / 

i 1 8*4! 7/7T 

1 // 

/ f. 





i if 

A $??# 

£ A ■ .. v'M 



<■-., ^r" '-0-iW. 





C' 1 '' 

-^'r^r^-^r*^^'^ 1 *' ' 


Fig- 4 Ot'Hosc/idnyastia cana n. g, ; v 'id n. sp A, dorsal view; B, dorsal 
scutum (x 500) ; C, l>s!p \\\ cloi-r-n i view (x 860) ; D. cliclicera (x 860) ; 
E, galral seta; T, dorsal seta (x^60); C Uu^us and nic:tabrGUs tIT (x)^50). 


Ascoschongastia'- ' uromys n. sp. 
Fig. 5A-F 

Description of Larvae — Shape oval. Length (miengorged) 3Q0ft width 
200/-/,. Scutum roughly rectangular with fairly well produced posterior margin. 
AM short and normally ciliated, AL and PL lanceolate or foliate, to ifii* wide. 


:-',• . 1,1:1 L ^ 

i Sill I 

1$ W « \ 

©if! @ 


f ir*s-# 


Ffe. 5 Asco^cho)!f,nisi(fi ueuuiys n. sp, A, dorsal view: R, ventral view; C\ 
dorsal scutum (x5G0); D, lip ol cheHcera (x860); E t galcal seta; F, dorsal 

,-mu ventral setae Cx3m)j, 

with longitudinal rows of strong dentures. Sensillae globose, apparently nude, with 
iheir buses only slightly in ad\anee of line of i'L. £,ycsj 2 -J- 2, on distinct ocular 
shields; posterior the smaller. Chelieerae with only the usual apical tricuspid 

< 3 ) Ewinpr (Proc. Biol Soc. Washington, 50, 19-!*"), p P 70-71) has recently shown 
that the genus Purascf'augasiia Worn, is sviionytnou.-: with Ncnschonfrisfia Giving" 1929 
(genotype- A", two 'Heaua), and for the remainder of the species o£ A"##£r/;xftt#&^% iiot 
ronspeoifje with aniCrictUia fo&ft proposed the name of yhco^chivif/aslui with A r . iiutfavt'Vs'ix 
Crater as genotype. The choice of this species a 7 tronotyyc is, however, somewhat un- 
fortunate, z$ it is v. rather aberrant species with PL distinctly oft' the scutum. 


cap. Galeal seiae nnde. Palpi fairly stout, with bifurcate libial daw. Setae on 
palpal femur, genu and tarsi apparently all nnde. Dorsal setae alt foliate or 
lanceolate with strong dentures (cf. fig. 5D), to 64 /t long and 16/* wide. 28 in 
number, and arranged 2.6.6,6.6.2. Ventraily with usual pair of branched setae on 
gnathosoma, a single ciliated seta on each coxa, a pair between coxae I and 
between coxae III. -and thereafter 5.6.6, anterior of anus, and of which the outer 
two Of J he second six approach Ihe dorsal setae in form.; -hen posterior of anus 
4 2 foliate and dentate as on dorsum; the anterior non-foliate ventral setae to 
20 fl lone;'. Ufesr: I 270 /i long, it 220'* fll 270 &} tarsi I and II with the usual 
dorsal sensory rod. III without any long nude seta. 

The Standard Hata derived from the t\pe and serai paudypes are as 
follows: . - . 

S';uidaid Th^oi-eHml jEtfreetVeil ColIF. ot 

Al^an Deviation tolp? Ran**.- Varialiun 

AW ... 50-fe1-74 4-92:bl-23 35-R-h.M 45-0-60-4 Sh7 

mV .... 69-5+0-71 2* 00 ±0-5-0 r>3'5— 75-5 60-0 75-0 2-0 

S!J - ,. I?*0±0-?0 G41d=(M5 14-8 2.3-2 18-0-21 -0 7-4 
ASB .... 21-0 No variation recorded 
PStt . 18-0 No vHrtauon recorded 
SI) .. . u f }'0 No variation recorded 

A-P ?A 'C±0-42 1 -19+0-30 18-0 25-2 21-0—24-0 5-5 

AM .... l<$-5±0«67 1-64+0-47 11-6-21-4 :5-C— 18-0 §4 

AL .... 49-5+0-67 1-^+0-47 44-6-54-4 48-0-5MJ 9^ 

PI. ,... a)-O±0-57 1-60+0-40 55-2 64-8 57 -Q— 63*0 2-7 
Seas.. 28-0 with bend 17/17. Only one tfeteniimir.ion 

Loc. — The tvpe and seven paratvpe? from Uromys lamiiujtoii from Dobo- 
dura, t?pw Guinea, 16 May 1944 (Xo. ol5, G. M. K.).' 

The type and two paratypes in the South Australian Museum, three para- 
types in the Rocky Mountain Laboratory, one paratypc each in the U.S. National 
Museum, and the British Museum (X. IT,). 

Remarks— This spfdes Glh- into the small crroup with foliate scuta! and 
dorsal se:ae comprising A. luisiata (Cater), inccitlloclti (Worn.), foliata 
(Gunther) and another specie^ at present unpublished. 

It is close to mcrullochi but aa: be distinguished by thj different number and 
arrangement of ihe dorsal and ventral 5etae, and by the t.cnlationa at the ,ciae, 
altbtui'.! i strcurj, bciup; nowhere so large as in lUCiullorhi. 

Ascoschong&Rti-'i eclrymipera n. sp. 
Fig. 6A-C 

Description of Larvae — Shape oval. "Length ( en^or-:-d ) to 460 /.*. width to 
550-pi ^Miliini roughly rectangular, rathrr 5 mall, and fairly long, with the 
posterior margin vcrv Shallow GhuG PL and lightly sinixm^; AM the shorto.&t* 
AL the lqrtjjftS; but only a Hide louder than PL, Sengn'ae g'ohose with sliort 
distinct eitiations ; the vensillae bases about midway b.'tweni AG and PL. Lyes 
2-| -2. on distinct ocular shields and -well separated Gem scuial mai^Gn; the 
posterior eyes very much the stmder. Cheliccrae with only the apical tricuspid 
cap. Galcal setae nude. Pa!j:i ;.touL, tibial claw trifurcate; on pit pal 
femur and genu branched, on tibia all three nude. T)S 34-.vS in number, arranged ,2(0 ) ( to 30 fr lon£ VgnttaJiy with a. pa'r of branched setae on 


gnatliosoma, a single ciliated seta on each coxa, a pair between co:;ae I and 
between coxae III, and thereafter ca. 6.6- to 26 ft long. Le-s ; I 21 o /.< 
long-, II ISOfi, III 226//.; taivd I an:! il with the usual dorsal sensory rod. 
If! wiibout auv lov.x nude seta. 

ic Standard Data idom the type and 12 parutvpe3 and 


other sped- 

jr. en 

r r p> 





55-1— ''.>/'• v 

4S>3bfc0^9 l-lfe:0-20 

Gl-5rh0-53 2- 12^=0-27 

20-4±0-29 3 -17+0-21 l-o 22-') 

21*0 No vaWalion recorded 

15-0 No variation recorded 

J6-0 Xo variation recorded 
2i-: + 0-27 l-49±0*2<> itf-fih^fl 

26-u=t0*3fj W5±0-26 22-2— 21-1! 

4o-5±Q-5-; 2'37±0-42 29-4— 53-0 

42-fefKV- I-^5rir0-26 22-2 4/4J 

27-0 vdtlt head 17/37. Mo rdtviiltm recorded 


Coeft. of 



57-0 — oC J >-0 


I£>0— 2!*tf 


40*1) 4» -o 

-I •/ 





jjw i 


^ t\ 

# / 2 ^ 

1 w 

Ld", 1 







Ascoschongastia tchyia'pcra n. sp. A, dorsal scutum (2x500) 
of cheliccra (xS60>; C pulp in cUrrsa! view (\-S60). 


/,i?;-. — The type and 12 paratypes from RcJiymipera cockcreUi from Dobu- 
dura.. New Guinea, 29 Nov. 1943 (No. 46, G. M. K.), and another paratvpe from 
same, host, locality and date (No. 48. G. M. K. ); also two other specimens from 
Rune and locality, 8 Dec. 1943 (No. 77. G. M. K.) 10 Dec. 1943 (No 81, 
G. 2d. K.). 

The type and four paratypes in the South Australian Museum, two para- 
types each in the U.S. Nat. Museum an I the British Museum (N. II.) ; remain- 
ing specimens in (he Rocky Mountain Laboratory. 

Rniuu'ks— In a new key to the species of /hcoschlhujasiia in preparation, 
rchynii^cra Trill run clown close to coorongense Hirst and innisfallensls Worn. 

and Heasp.. but is easily distinguished by the scutum and the number and arrmige- 
mem of the DS. 


ByT. Harvey Johnston and S J.Edmonds 


Parasites of this species were found in the fish Callionymus calauropomus, caught in St. Vincent 
Gulf, South Australia. Five of six fish examined contained this parasite, and in four of them they 
were abundant. The intestines of all these fish contained crustacean material, especially amphipods 
and cyprids. The collections examined contained both males and females, the latter being slightly 
longer and broader than the males. In all the specimens examined the proboscis was protruded, but 
in no case was the copulatory bursa of the male everted. 


No. 5 

io T. II \ryly JoiiK^TON and S. )- ! ; . r jMC\*DS, Department of Zoology. 

University of Adsfljdfe 

[Real 10 October 1946] 

IJvPOKCHiMORiiYNCiiUE Ai.Auor-is; Yamaguti 19,39 
Pfife 1-9) 

Parasites of ffus species were. found iu tlie hsh CuKiOiiyjnus cirahroynuts* 
caught in St. Vincent Gulf. Sovtk Austrara, I r ive of six fish examine:! con; 
iame:i this parasite, and in four of iucrti tin- y were abuu-lant. The iiileslines at 
all ikc c c hsh contained crustacean material, e-specia'y amphipods and eyprids. 
The collections examined contained both males and females, (lie kilter being 
slightly longer and broader than the ira''-?, Tn all rhe specimens examined the 
probc-cis was prclruded, hut In no case was the copulatory bursa of the male 

The T en£th of ihe male is 1 • r] to 2*7 mm., and the female 1*6 to 2*8 mm. 
The maximum wi.hh o. the male is 0*45 to 0*70 mm,, and the female. Q'50 to 
- 91 mm. The body in both se>:c-. i$ curved vent rally anu devoid of spines. The 
proboscis U globular 1o spherical in shape and is attached 10 the trunk veutro- 
Rrmmally (Eg. 1). Tkti proboscis oi the male is 0*10 to 0T4 mm. lon-f and 
0-10 to 0-15 mm. iu ks u dde>t pail, live corresponding measurements m the 
feina'e are - 1 2 to (hi 6 mm. Tie neck portion of the oroboscis is A r ery short. 
The proboscis is armed with 25 hooks arranged in ten longitudinal roves, consist- 
ing of hye rows each of three hooks alternating \\\ih (he rows each of two hooks 
(fur, 4. 5_>. The Lengths of the hooks, measured along the curve from the point 
of er-tru-sion to the tip of the hook, are shown :n the following table: — 

MiJc Anferjot >firU!lc r/ostmor 

£uw oi three 78-101 t u oi~oS p 21-2^ jU 

"Rovv of two /l-89 ( ,,t, 25-32^ 

Female A^tc-W Miiklle Pij^ltriur 

Row of three ,... .„. S^-UO^ JM? fj, 23-32 ^ 

Row of two ,.. ,.- 7t-90^ ^J~35 ^ 

The proboscis thcarh is bu'b-bke and in the male measu.e>> 0T2 to 0"?.2 mm. 
long and 0*11 Jo 0T4 mm. in in wides- pa^t. The eotrospondnufj; measurements 
fri the female are 0T2 to 0'20 mm. and 0T1 to 0T6 mnt. The sheath, which is 
inserted at the base of the proboscis, \:^ double- walled. The maximum thickness 
of each layer in both sexes; 5s. ID to 15 p* A spindle-shaped gangoon is sdoated 
at the posterior end of the sheath, aud the rei'macula arise from the bide walls at 
about thi> level. A strongly -developed rotrae;or is present in boib so;ces. The 
lemnisci arc shorl, stout and cylindrical, and contain a well-developed lacunar 
system. The hypodermis is thick and the lacunae of the body wall anastomose 

MuL' system — There are two spherical to oval-iikapcd testes placed one 
behind ike other hut usually pressed close together. They lie in the anterior part 
of the worm. The anterior testis h slightly larger than the posterior, the dimen- 
sions of iltc former being l>21 to pf"3S mm. long and 0-20 to 0-26 mm. wide; 
arid of the posterior 0'20 to 0-^2 mm. Ion# mid 0'18 to 0'25 mm. wide. Two 
vasa ctTerentia. unite near the anterior end of the Saeftlip;cn*s pouch, to form a 
common duct, which in mot-t specimens is swollen at its base to form a seminal 
vesicle. Thir> terminates in a penis which projects into the atrium oi the bursa 

Trana.Ro>. Soc R. Ajst., 71 CH, 23 July 19 M 


and which is enclosed in a capsule or genital papilla. There are six cement gland? 
which are elliptical to pyriform. In most cases they lie pressed closely together. 
The ducts of the six glands unite to form two lateral ducts, which join at their 
bases to form a U-shaped ccmenl reservoir. Two well-developed diverticula 
project anteriorly from the bursa. The genital pore is terminal. 

Fig. l- ( ) — Hyl'Oftvuaothyiwftits ohwrf^ 


J : cwalc system — The general anatomy of the female system is shown in 
fig, 6. la a typical specimen lbe uterine bell 35 0*15 nrm. te'njg and is separated 
From the uterus by a narrow constriction, 0*12 111111. in length. The uterus in the 
same specimen is 0-35 mm. long" and 0*07 mm. wide at the anterior end. Some 
of the female specimens contain floating; ovaries, while the others have both 
ovaries and eggfc Mature eggs, when mounted in methyl salicylate, measure 
50 to 54 w lUHff and '3 to 16 {^ wide, The polar extrusions of the middle shell are. 
well developed. A nuinher of females bear a copulatory cap at their posterior 

Systematic position — We consider that this parasite belongs to the species 
llypaecliiiwrhxui'hus alfteopis, described by Yamaguti (1939, 325),, from a 
Japanese fish. Our measurement? agree very closely with those given by him. 
The hooks of our specimens, however, seem slightly longer, and the testes some- 
what smaller than in his materia]. We regard Hypocctiinorliynchus as a valid 
genus of the family bldnnorhync'iidae. The form of the proboscis, as well as 
ihe shape, number and arrangement of its hooks, arc suggestive of those of 
A r cacchinorJiy nchits, but the characters of the cement glands are quile dirlerent 

Although the parasites described by Yamagult were taken from Alacops 
piinthiis, he stated that a single immature female specimen \va^ collected from 
Cdlliou ymus altn^Hs. 

Pararhadinorhynchus mugilis n. gem, n. sp. 
CFis. 10-22) 

This species occurs in (lie mullet, Muyil ci'plntliis. Five of six fish examined 
from l\m Wilhmgn in Mnrrh, 193^, were parasitised, in one case heavily. The 
intestine of all Lhe fish contained much plant debris with occasional molluscs and 
small cruslnceans. Two other fish taken at American River, Kangaroo J stand > 
by Mr, Tf, M, Cooper in January. 1915. also contained the same species of parasite 
in considerable numbers. In the stomach oi these fish were gastropods and 
numerous crustaceans (prawns, anrphipocls and copepods;. Both male and 
female specimens were collected. The worms are long ar.d cylindrical, the female 
being longer and slightly broader than the male. Both sexes are devoid of body 
spines. The length of the male ranges from 3*1 to 11 '4 mm.,, aud the iniuirmmi 
width from 0*23 to 061 mm. The females arc from 3":> lo 19*2 mm, long and 
from 0*22 to 0*69 mm. wide. The posterior half or third of the female is twisted 
in most cases into two or three spiral-like convolutions- -i'iig. 19). Tins may be 
due to the fixing processes, All measurement^ were marie on animals cleared in 
methyl salicylate. 

Although the collection consists of a considerable mmiber of both sexes, in 
only two is the proboscis fully It is therefore diiuenli to give a range 
of values for the length and breadth of tltaf organ. Tie.* proboscis in most adult 
specimens is about 0*9 imu. long aud Ch2 mur wide in the broadest part. It 
appears to taper 5 lightly towards the base. It bears IS longuudiual rows of hooks. 
most iji which are firmly attached hy rooting processes to the cuticle. Each row 
consists of I'G to 17 hooks. Tin 1 form of ihe proboscis is shown in fflf. 10, and 


1, male; 2, female; 3. proIio?ri^; -1, 5. row •? of hooks: lemnlc nrflaus; 7. e^; S, T\S r final*?, 

tluouqh ccaicut j^'brit.!?; 9. T-S. male, through cement chiefs, 
b, turr'a, I_ir ( brnin; Inv, borly \\:i\\; vc, conidatai'y cap; H, a ipent tfuet : ci. coagulated 
body flu if I; ce;< cement gland; et, cement reservoir; t'j, ejacaknory duel: g, ii.aii;;lion : 
fto. genital opening: #:». ^emtat papilla; 1. lemniscus.; la. lacuna'; I^_, ligament; Ini, lohf.o'- 

tiuhiial miHrle; lo, tltcrnl opening o| nO'rinr brll; n, nucleus; p<\ proboscis; pi. proboscis 

theiit; tni irtr:u:tnr muscle; S,i. S/i eff ti-^onV pouch: Ffih, sphnrier; r, icste*: u. uterus; 

lih, uterine be'l: vb T vafc/ntsl bulb; vd, vu.^ deferens; vr.. ve.dcubi >e_i:m;dis, 


■L'i*. 10-17 

Ctrnr^r ;,\i i^ VEWftfJf 


1 -rnLo^cis o: m:i!e; 11, hoi 

fivm anterior, mkKk- and posterior regions of proboscis; 13 posterior end o( 

i:i; ly: l3i '1 •& anterior mil of male; 14, T.S. Mirougli rc:?io:i of ctment glands; 

15, I'.S. t!ilOfiisrh cement chirrs; l'u T.S. thvoMtfli re;;':on of cj;ic;.lutory duct; 

17, T.S. uejon of R&KjrHrtfi of 


tlu- siVe nr.d form of the hooks in fig, 11. There is a slight neck region. The 
proboscis sheath is double-walled and measures from 0'6J to 1*3 mm. }i:n-^ and 
from 0*12 10 0-20 mm. wide. Tlic thickness of each wall i| about 0-02 nirr. A 
bruin is situated towards the bate of the probose-s sheath. 

The lacunar system of the body wall show? two well -developed S^dgittiSiii^] 
lacunae, from wlrch nuastomosmjC;" channels aHre (fiir. 22 ). Namerov.s -mall 
nuclei are found in the body waT 

The lemnisci are about 0*8 mm. Iftng an 1 p-sleflH tifttelty p.s far r\i- the 
posicrior portion of the proboscis gk-istfk. Transverse -.unions c.f the knmisei 
show that they arc fiat m& two :a.cra; caim's an] a ''ember of lar^v rvudej 
are al^o present in these structures ( fe,r, la). 

Miilv system — There arc two elongate rcslcs which iju close, together, one 
behind 'he other, in the posterior bait en* t'.rrd of ihe au'niuh The anterior ;_esbs 
measures from 0-28 to 1*1 mm long aftd from (HVi to 2*1 mm. wide, and tbe 
posterior 0*27 to 1*1 mm. long ami Q-flS to 0-23 mm. ivtfl'e. There st-e two long, 
narrow cement glands which rrm.^e fa to tog£*fe from 0'45 to 2*5 nv.n.. and iu most 
tpeeimuw they atv swollen posteriorly. Tbe tiwei^ from these glands f'-rre two 
long cement reservoirs which usually ;uv eonrtrtelc-d in cue or twer places .ovards 
their posterior extremities. A long Sactfchjen's* pouch lies between *he t\vo 
cement duets and reaches forward as far as the distal ends of tbe, cement ei.inos. 
The va-5 deferens swehs slightly towards hs posterior pirt to form a seminal 
vehicle. There is nn ejacukitory duet and u well-developed bursa which bears 
ray-. in none of $w» specimens examined was *bc bursa evened. The male 
aperture is terminal and is surrotiuded by tnnre-ous coll.. probably constituting* 
a ganglion. 

ficmah* sysh'iu— Tbe structure and arrangement of the female -y^um is 
shown h* lb;. 20. The uterine beil is about 0*20 mm. Jo- g. The uterus proper 
in mature specimens ranges from 1*1 to 1* 1 mm. in length. The genital opening 
is terminal and the ganglionic complex which surrounds it is about IH5 man !oug. 

Mature eggs, measured in 70% aieohob range from 56 to 62 /i long and 14 to 
\$H wide and bear polar prolongations oi the middle -died. 

Syxter.iatic fiosiiinr — This species doe:> not fit. very well into Van (..'leave's 
conception ( 1 r >2^ ; 1Q1Q) of the "Rhadmoiii'.r.dndae, hut it r-escmhic? mot of the 
members of that family in the form of the proboscrs, the thane at the hook--, the 
double-walled proboscis sheath, tbe fofl£ ( (ubular cement ghinds, and the feci thai 
its ho-. U a fish. Tt: (litters from l he known gvngrci of uH family in the lark of 
body spines, though one such genus. Lrplorhyuchmdrs, has already been admitted.. 
We propose for the reception of this species a new g'enus, Payarltadhwrliy.i kits, 
with the following characters: — Rhadiuorbynehmae ; body elongate* cylindrical : 
proboscis- long, wldi numerous hook* ; proboscis: sheadi dumle-uallwl with pro- 
boscis predion towards the posterior end: body devoid gE spines; cement ejand-, 
two, letjg. tubu'ur, swollen sii^ldly towards the posteri:-:* end; numerous small 
nuclei tn body wal! ; genital Lvan^uon wdl deve'opef!. in tish. Type. i\[ra- 
rluufiiwrkyiicints iinifjtfis. Types: have been deposited in the South Ausisahan 
Aluscuin, Adelaide. 

l^uAnj^niuiv^CHUS pr.i.stts (Jiuoolpht 1802) 
<>>. 23-25) 
( )uc immai lire female of this species v.a 1 -; found in I he iutestlne of the 
southern tunny, Tlwnnus maccoyi, caught off the Semaphore in Sr. Vincent Gali, 
South Ausireiia. Tbe worm was ioi\£ and tubular, its length being 17T mm. 
and its maximum breredlh 0-60 mm. The proboscis, v, hah was fully retracted, 
was 1*9 mm. long* and bore many hook:-;. Tiie proboscis sb:ath was double-wailed 



Tig. VS 22— Fanii'hadinorhytickiis inugiHs: IS, male; 19, female; 20, feni^c 

organs; 21, egg; 22, part of lacunar ^ stem. 

Flgi 23-2.7 — Rhailiiwrhyiidnis prist's : 2}., anterior end of female ; 24, body 

hooks; 25, egg. 


and 2*8 mm. long. Two lemnisci extended back as far as the posterior part of 
the proboscis sheath. The extreme anterior part of the body bore two sets of 
hooks, one group surrounding that part of the body adjacent to the proboscis, 
and the other group lying on the ventral side of the worm. The shape of the 
hooks is shown in rig. 24. The specimen was filled with unripe eggs which pre- 
vented the examination of the female complex. The largest of the egg* were 
0'062 mm. long and 0*012 mm. broad. Three shells were seen, the middle bear- 
ing well-developed polar prolongations (rig, 25). 

The specimen agreed in all essential details with the figures published by 
Liihe (1911, 44-46 3 fig. 58-63) and Meyer (1932., 47-48, fig. 23-25). The latter 
mentioned several kinds of fish as hosts of the parasite The species is now 
recorded for the first time from Australasian waters. 


Luhe, M. 1911 Acamhocephaleu. Die Siisswasser fauna Deutschlands. Heft 

Meyer, A. 1932 Acanthoeephala. Broim's Klassen und Ordnungen des Tier- 
reichs, Bd. 4, 2 Abt, 2 Buch, 40-62 

Van Cleave, H. J. 1923 Key to Genera of Acanthocephala. Trans. Airier. 
Micro. Soc, 40, 184-191 

Van Cleave, H. J. 1940 A Reconsideration of the Family Rhadinorhyiichi- 
dae. Jour. Parasitol., (1), 75-84 

YamAguti, S. 1939 Studies of the Helminth Fauna of Japan, part 29, Acantho- 
cephala ii, Jap. Jour. Zool., 8, (3), 317-351 



By J. M. Black 

Acacia euthycarpa desc. Em. Frutex magnus glaber, ramulis teretibus; phyllodia usque ad 8 cm. 
longa, 1 mm. crassa, teretia et 4-nervia, vel leviter compressa et in quoque latere duobis nervis 
additiciis inconspicuis instructa; capitula gemina vel pauca in racemis brevibus efoliatis; flores ut in 
A. calamiformi; legumen subplanum marginibus strictum, 8-12 cm. longum, 6-8 mm. latum; semina 
longitudinalia funiculo conduplicato cincta. 

Southern district to Flinders Range; Murray lands. Differs from A. calamiformis chiefly in 
the straight-edged margin of the pods, which are not or very slightly narrowed between the seeds. 
This is a more complete description than was given in these Transactions, 1945, p. 310. 



No. 44 

By J. gl. Ih.ACK 

[Read 10 October 1916 j 

Acacia euthycarpa de^c. em. Frulex magnus glaber, ranutHs terettbus; 
phyllodia usque ml 8 cm. longa. 1 mm. crassa, teretia et 4-nervia, \xi levitcr eoiu- 
pressa ct in qnoque latere ritiobis newts additions inconspicuid instructs; capittua 
gemiua ve! pattca in raeeud? brevihus efolialis; florcs ut in aft culantifortnt ; 
leguinen suhplamun margimhus ^UictUnI* 9J2 '-^i. l^i'gttni, 6-8 mm. latum ; seinirm 
longUudinaiia fuuienlu conduplicaio cinrta. 

' Southern district to Flinders Range ; Murray land*. Differs from A. catcwi- 
formh chiefly in the straight-edged margin of the pud-, which are nor a} very 
slightly narrated between the seed?. Thi- is a more complete description than 
was given in these Transactions, 194-5, p. 310. 

Acacia pingutfoKa nov. sp. Frutcx glaber. rauudis Icretibiis; phyllodia 
carnen tereiia erccia vel recurvata 24 cm. tonga 2-?, mm. cras-;sa obscure circa 
12-nervia mucrcne brevi rluro terminata ad basin artieulata; peduneuli solitarii 
vel gemhii 5-10 nun. longi ; capittda circa 25-flora; scpala 5 Hneari-spathutata 
eiliala dinridio hreviora quam pera'a; bvaeieulae clavuHfornies; legumen e:rea 
3 cm. lomrum 2 nun. latum contort uni & super semina longit.udinalia convexum ; 
semen nigrum; funiculus hluornns, in arillum crassum album uuadruplicein 

Finnis,. River, near Lake Alexandria: near Port Lincoln. Near A Bynocajw., 
Benth., and A. MeiicAii, but is net viscid and has thicker phyllodes with inure 
numerous nerves. 

Differs from A. sulcata R. Fir. in the rFshy, much thicker ami longer 
phyllodes. and the monilifonn pod; from A. ganophylla Benth. by the phvilodes 
not tetragouous or incurved aud by the pod which in A. gonophylia is flat, straight 
and not twisted. 

Acacia ccrcnuli's nov. sp. Hamuli subanguh'ti; phyllodia glabra, ovato- 
ublonga vd lato-ianeeohta, coriacea, basin versus angustata, 2M$ cm. longa, 
6-15 mm. lata, obtusa* oblique l-ucrvia, pemiivenia, margiullms uemfonnibns; 
stipulae praecipes* spinosae : caphula parva, g-r.bosa., 15-20 in cpuxp-te racemo 
axiilari dhposita, 20-25 flora, tenucs. pubeudi, peduneuli circa K) mm. tpilgf, 
solitarii. geinini vel terni, raccmos phyllodiis st.tbaequulibus formantes ; scpala 5, 
i.mean-spathu'ata* 1 ftiftx 1or,ga. aptee ptlosa. petahs dupio breviora; ovarium 
glabrum, brcvissimc stipiiattmi; iegumen ignotum. 

Crown Point, River Finke, Central Australia. The exact re'kticm to 
A. Victoria?, Benth., cannot be known until the. pod is found. 

Dsviesia nuduia nov. sp. Frutcx stnetu.s glaber 1-2 m. ahus; ranudi 1ongi 
rigidi su-catt-tereies apiee pungentes p^rkiue aphylli, cjiiandoque paueis loins 
brevibas (3-9 mm. bugis) sub u' at is spiniformibus divaricatis pungentibus 
praediii: folia etun ranuills roi^inuu; dote-; 3-12 in racemis solitatirs vcl con- 
j.ire. f iuTds i-2^ cm. lotrns fere oninilms lateralibus; pedicelli capiliarcs patentes 
5-12 mm, lon^i; calyx 4 nun. luilgUS turhinatu? in pedTccHiun pattiutim fasii.L;atum. 
i/CTjtmus brevibus acuus, duobus superioribus hrevioribus appro .\-inmnsr{ue : caiiua 
obtupa; lerjumen irmriatunun triangulares 

Berri (River Murra.}*). Perhaps the satire as D. briz-ifofia,, Lindh var. 
ephedra ides, l^enth... oi Western Australia. Differs irom D. hrevifnlia in i^ 

JiattS Koy. Soo. 3. Aust,, 71 (I). 25 July 1^7 


almost leafless character and in its much longer racemes with capillary pedicels 
usually two to three times as long as the calyx. 


Grevillea umhellifera now sp. Funiculus 60-100 cm. altus, ramis validis, 
asperis, fuscis, pubescentlbus ; folia erecta, angustc linear ia, rigida, apice pungen- 
tia, 5-10 cm. longa, 2 mm. lata, supra glabra, intra puberuhi et. bisulcataj integra 
aut pauca summa bi- vel tripartita, omnia rigide hreviSerque petiolata; florcs 
pauci usque 20, pedicellata, in urubellis vel capittllis fere sessilibus, plerumque 
axillaribus, congregati ; periaiithium extus roseurn, dense puberuluni, interne sub- 
turgidnm, iutus saturate rubrum glabrumque, tubo 10-12 mm. longo ; ovarium 
glabrum ; gynopliorus ruber, in feme perianthii basi adnatus ; glandula nulla; 
.stylus glaber, ruber, 9 mm. longus, disco lateral!; pedicellus puberulus, 4-5 mm. 
longus; fructus globosulus, niger, laevis, circa 12 mm. diametro. 

Koonibha, near Fowler Bay, Sept, -Oct., J. B. Cleland. 

Near the Western Australian C. brachystachya Meisn. in leaves and in- 
florescence, but differs in a few of the upper leaves divided inro two or three 
linear segments, the flowers larger and the lower part of the gynophore adnatc 
to the base of the perianth. 


By T. Harvey Johnston and Patricia M. Mawson 


Most of the material dealt with in this paper has been collected over a period of very many years by 
the senior author in New South Wales, Victoria, south-eastern Queensland, Central Australia and 
South Australia. We are indebted to Professor J. B. Cleland for some of the specimens from New 
South Wales, and to the late Dr. T. L. Bancroft for those from Eidsvold, Burnett River, Queensland. 
Acknowledgment is made of assistance from the Commonwealth Research Grant to the University 
of Adelaide. Types of the new species will be deposited in the South Australian Museum. 



By T t Harvey Joiintsto.v and Patricia M. Mawsox, 
Department of Zoology, University of. Adelaide 

[Read 10 October 1946] 

Most of the material dealt with in this paper has hcen collected over a period 
of very many years by the senior author in New South Wales, Victoria, soulh- 
eastern Queensland, Central Australia and South Australia. We are indebted to 
Professor j. B. CI eland for some of the specimens from New South Wales, and 
to the late Dr. T. L. Bancroft for those from Eidsvold, Burnett River, Queens- 
land, Acknowledgment is made of assistance from the Commonwealth Research 
Grant to the University of Adelaide. Type* of the new species will be deposited 
in the South Australian Museum, 


Varan OS vakius Shaw. Physafoptcra anlarctica Linst. (Burnett R., Queens- 
land). Fhysoloptera confusa J. and M. (S.E. Queensland). Qphidascaris 
vaytini n. sp, uS.E. Queensland). 

Varan us belu Dum, and Bibr. (Probably only a northern variety of V. varitts). 
Physdloptera confnsa J. and M. (Burnett Rj- 

Varanus GGtxm fjrav. PhysalopU'ra antardka Linst. (Burnett R., Oneensland; 
Victoria). P, confnsa J. and M. (S.E. Queensland; Victoria:. 

Varanus oh.iaxteus Gray. Physaloptaa confnsa J. and M. (Central Australia). 

Varanus. im:xctatus var. orikntalis Fry (Eidsvold, Burnett R.). Physahp- 
tcra sp. 

Timqua scincoides Shaw. Pimtmoncnta tlliquae Johnston (New South Wales). 
Phvsaloptera aniarcika Linst. (New South Wales; S.E. Queensland). 
Pharyncjodon anstrafc J. and M. (Burnett R., Queensland j. Pharymjodon 
tlliquae Baylis (Burnett R.). 

Til.ioua nigroi.utea Gray, Physaloptera aniarctka Linst. Physaloptera sp,, 
zrA Pharyngodon attstrak J. and M. (all from Katoomba, New South Wales). 

Teactiysaurljs ulgosus Cray. Thchmdros iracfiysauri n. sp. (Adelaide, South 

Egernja cunntx«:iAmt Gray. Pharyngodon tUiquac Baylis (Bathur-t, New 
South Wales). 

Eukhnia STf;jor.-vr-\ Peters. Pharyngodan tlUqitac Bayb's (Sydney and Kendall, 
New South Wales). 

Egkrnja whit™ Lacep. Pharrtvjodon tUiquac Baylis (Sydney, New South 
Wales L 

Eoerma dahli Boulenger. Flurry ngo'don tiUquve Baylis. AmphibivphUns 
cgcrnktc n. sp. (Both from Mnsgravc Ranges, Central Australia.) 

Lycosoma quoth D. and B. Physaloptcra sp. (Lower Plawkesbury River, New 
South Wales). 

Oepura kogusta Boulenger. Physaloptcra sp. ; Parathclandros ocdurr^ m sp. 
(both from West Burleigh, S.E, Queensland), 

Trans. Roy. Soa S. Aufct., 71 U)» 25 July 194/ 


Amphibiophiius egemiae n. sp. 
(Fig. 1-3) 

Two males and one female trichostrongylid worms were taken from the 
intestine oF Egcrma dahli, Ernabella, Musgravc Ranges, Central Australia. The 
males are 3-9-4*4 mm. long, artel the female 8'3 mm. The body is spirally coiled, 
and narrow lateral alae are present. The mouth leads to a small buccal cavity in 
which is one tooth. The oesophagus. \1 mm. long in :he male, is somewhat wider 
in its posterior third, and is surrounded by the nerve ring ]w\ be lore this level. 

The vnlva lies near the posterior end/-2 mm. in front of the anus, The tail 
is *07 mm. long. The egg nearest the vulva is oval and measures 65 /a x 30/*,. 

The bursa is rightly folded in both male specimens available, but the lateral 
lobes appear to be symmetrical. The dorsal lobe is hardly demarcated from the 
lateral lobes. The arrangement of the rays is indicated :n the accompanying 

Fig. 1-3, Aml'k'l/iothikis iVH-rtr'ac; 1, ticad; 2, male i-atf; 3. posterior end of female 
4-6, Opfnddscnns varani; 4, 5, lieadi 0, mule tail. Fig; 3, 4, 5 and 6 to same scale. 

%. (fie;. 21. T!:e external dorsal ray appears to aviso from beside the base of 

the dorsal, ralher than from the latter. 

each of 

with tl 

long is present. 

The species appears to fls to be the nearest to those of the genus 4>Ulf*M 
biopliilus, differing from them, however, in having four, instead of the :h 
terminal processes on each branch of the dorsal ray. 

g. 21. T!:e external dorsal ray appears to arise from beside the base of 
rsal. rather than from the latter. The dorsal gives riflfl to Lwo branches, 
f which has ftijjt tenninal digits. The spicules are simple, '2 mm. m length, 
lie proximal half wider than the distal A gtibernaathun. -O45-*05 mm. 


Ophidascaris vprani n. $p\ 

(Ffcr. 4-6) 

A single mn!e worm belonging to the germs Qflvdasntris was taken from 
Varavus writs* from Toowoomba, Queensland. It measures 7 mm. in length. 
The very short interlabia are separated from the lips by deep grooves, and both 


interlabia and grooves are bordered by a wide continuous ciiticiilar flange. The 
lips have a very narrow lateral flange, bearing a line clcmigerous rid^e. The: 
oesophagus is 4*8 mm. long, The position of the nerve ring is not clear. The 
posterior end narrows suddenly al the level of the arms- The tail, *15 mm, long-, 
bears two pairs of lateral and two pairs of sub-median papillae. There are a pair 
ol! double- headed pannhte. and about 30 pairs of small pre-anal papillae. 
The presence of median pre- and postanal papillae is doubtful The alate 
spicules are '9 mm. long; thcT length lk"*<g much shorter then any hitherto 
recorded for species of Ophfth$fttyi$. In adoirion to this characteristic ltt« 
present specimen differs from the oilier species o[ the W?&p& recorded as occurring 
in Australia. O. filcna (Duj.) and 0. pynltus J. and M. f m the length of the 
imeriabia and in the number a! caudal papillae, 

PrrvsAr-nFTHRA Antarctica Linst. 

This species has h-Btffi taken in larg-c numbers from Til<qua sciiicoulcs { 'Xcw 
South Wales and S.E. Queensland); Vanr,nis mum (lUiriiett River, Queens- 
land; and Victoria) ; T t -;n)jrolutra (Kaiootnba, New South Wales), and 
V. tvr ins (Burnett River, Queensland). 

A very great variation in size has been observed, mature female specimens 
in some collections being only about 2 cm. long, in others reaching 5 cm. 


This species,. Which was originally recorded by us (1942) from the t%er 
snake, Nolrchis sattnius, is very dose lo P. antardica Uiist., as redescribed by 
Irwin-Smirh (1922, 57; 1922.233) from material from TWijiut qfy^ and Varanus 
vorius (1922. 419). It was suggested by us thai P, confnsa might be restricted 
to snakes. This, however, does not seem to lie the case, since among a large 
number of Physalapterids now examined from Verauifs spp. some are determined 
as P. cniwrctiea and some as P. confusa. The latter nematode is now recorded 
from l-'anuius f/otfldi, Calnuudra, South Queensland, and Victoria; W' m vorins. 
Toowoomha and West Kurle-'^h. Queensland; V. hvlli, liirlsvold, Burnett River, 
Queensland; and l~. (jifiaHiruSj Ilermannsburg;, Central Australia. 

PKVS\I,ni'Tti-:;\ Sp, 

In some collection? there were Physaiopterid worms in an immature condition, 
or too poorly preserved for sa:b*f actory examination, so that the species could 
net be determined. Such material was obserwd in collections from Tiliqi'u 
nhjrrlvhra (Katoomba. New SfiUtu Wales); Patflkns {"unctafvs var, oai'n/alis 
(Kid; void, IHirnett River) ; L};;o$o;itu qiuiyi (Jlawkesbury R., Xew South 
Wales); awl Ocdura robttsta (West Bnr!eie;h. Queensland). 

P-\-i:r_\iONr,M_'\ TiLiru;Ar Johnston 
Many specimens of this parasite of the lungs were examined from the type 
host. PiUqaa scincoldes, from Sydney, As in the original materia], females only 
were present. 

Thelandrcs trachysauri n. sp. 
(Im-. 7-8.) 

Prom Truchysanrus mr/osNS, Adelaide. Males up to 2 mm.. Females to 
3*5 mm. Vestibule praciically absent. Oesophagus "55 mm. long in the male, 
'6 mm. in the female. Excretory pore post-ncsophagea). Nerve ring about '2 mm. 
from the anterior end in the male. The vulva is 1'4 mm. from the po?rerior end 
of the body in a female 3'2 mm. long, i.e., just posterior to the mid-body. Eggs 
are thin-shelled. 51 ^ x 100 yt. The female tai! lapers gradually to a ime point, 
:md measures *4 mm. in terfcmi, 


The male tail, or dorsal spine, is "1 mm. feflg and bears one median papilla 
at about irs juid-Iength- 'There are a pair 01 prc-anal and a pair orf poM anal 
papillae, and a median post-anal prominence wheh is a true papilla. The spicule 
is -1 mm. lon^. The species closely resembles T. hariur.a L and M„ from which 
ii is distinguished by the greater length of the spicule t.nd the different shape 
of the ?&k* 

B%. 7-3- Thrfandr^s (rarhysauri; 7, male; 8. female. 9-13. l } arcit/ii'la;u(rp$ 
ocd'irac; 9, head; 10. posterior ax! of female: 11, posterior end of male; 
12, mule tail, ventral view; 1.5,^rior tixl of female. 0%, 9, 10 ant! J2 to 

same se«lc: iff; 

and 13 to £* 

ane sea 


Parathelandros oedurae n. sp. 
flfigi 9-13) 
From Ocdura robusta, from West Burleigh, South-east Queensland, Collec- 
tions were made from several specimens of this host species. Males up to XtB mm. 
lo,H3 and "24 mm. wide; females 6 nun. long and \YZ mm. wide, inclusive of the 
tail spike and lateral alae in both cases. Body tapering markedly toward? head. 
Lateral alae present from about posterior end ot oesophagus to level of anus, 


and relatively very much wider in male than in female. Posterior end in both 
sexes bears a long, tapering spincd spike. In the female ihe posterior end bears, 
in addition, rhree stout backwardly directed "horns" which are .strongly cuticular- 
ised extensions of the body v^all. but are uol so long" as the tail spike. Oesophagus 
"3 mm. long in the male, *6 mm. in the female, its anterior part cylindrical, and 
ending in a large spherical hu'.b '13 mm. in diameter in the female. In the adult 
female the rectum is very narrow and quite insignificant Nerve fiag '13 mm. 
in male and '15 mm. ill female from the anterior end of the body. Excretory 
pore post-oesophageal, ju=t in front of vulva in (lie female. 

Male; Body tapers abruptly at level of cloaca to end in spike v nun. long, 
on which are a pair of papillae and about 12-15 small opines (jig. 11). A pair 
of pre-a:;?.: and a pair Crf po^t-c.nal papiika:* arc present. One spicule. "II mm. long, 
and a gubernaeulum about '03 mm. lone;, have been observed; the- presence of a 
second spicule is probable^ b;:t was not seeti. perhaps owing to the concentration 
of tissues in this region. 

In the female thft tail spine is about "9 mm. )nug. and bears abont 16 small 
spines. The horn-like processes nt the pnst< 'ricr end of the bodv projrer are 
■13 mm. long (fig. 10). Cterus narrow, but very long and much twisted, its loops 
extending behind the termination of the intestine. Eggs 142 m ?c 43 m, Vulva at 
level of oesophageal bulb or just posl-oesopbageal. 

The species differs very Jktle from 1 J . minlis Cbitwood, the distinguishing 
features in Ihe male of the present species being the absence of a median post -anal 
papilla, the presence of tail sivnes, and the presence of a spicule; and in eIic 
female, the form of the posterior end of the liody mid ffig size of the eggs. 

PtiAkYXGOPox Dicsiug 1861 

The ge'ius PJtaryngotfon has been rrcordcj several times from Australian 
h&firSj four species being named from reptiles. The&e species ai'e not. all inemded 
in the vahialr.e analysis of. arid key to. the gams published by Spaul {1920), since 
P. I:ii:dfri Thapar 1025 was the only one of them described prior to 1926. Fol- 
lowing that key. all the recorded Australian species fall into the group "Aa*\ 
that is, species in which caudal alae of the male include the pair of post-anal 
papillae, and in which the adanai pair of papillae are forked, \Yc have further 
differentiated this group as follows:* — 

1. Fcimik- tai: opined. P, bi-rUp-.a J. ami M. 
Ffcsjttla toil naked. 3 

2. Si^ke of male tai'. Uvu to tlircc times length of Imr^a, I\ iJnnuii' Hi\\\i> 
Spike longer thari, hti! col twice length of, bum. P. h'trJL'i Thapar 
Spike sl.orter than length of bur u :r. P. misi-'^Ji' J. ;aul M; 

Oliver differences between these species concern the position of the excretory 
pore ar.d ihe s : ^e of the, eggs, II ts remarkable that three of ihcm should have bceu 
recorded from the sam^ bo^t species, TlUqita schicoidcs. As far a> the informa- 
tion at present available 15 concerned, ?U these species appear valid, 

Pnw-wirox tiliov-U; Haybs 1930 

This ^pt-'des was originally described from Ttliqtui srincoides : it is now 
recorded from thru host from Kidsvoki. Queensland ; lif/rrnia* (Sydney); 
/I. siriotaia (Kendall, New South Wales); E. iunrnaglu-'Ml (Pa'dinrsL, New South 
WaL-'O ; and II. dxlui (Aiu^rave Ranges, Central Australia). Pharynrjodon >p.> 
described by Thapar (1925) from female specimen ^ front li. r;t:itihiffhuvii agrees 
with the description given of P. filioum. urn! aw. now be placed in the synonymy 
ci the latter. 


Pharykgodon australe J. and M. 

Originally described by us (1942) from Tiliqua scincoidcs, from New South 
Wales, this species is now recorded from the same host species from Eidsvold, 
Queensland; and from Tiliqua nigrohttea, from Katoomba, New South Wales. 

Baylis, H. A. 1930 A.M.N.H., (10), 5, 364 

Ikwin-Smitii, V. 1922 Proc. Linn. Soc. N.S.W., 47, 53-62; 232-244; 415-427 
Johnston. T. II. , and Mawson, P. M. 1941 Rec. S. Aust. Mus., 7, (1), 

Johnston, T. H., and Mawson, P. M. 1942 Rec. Aust. Mus., 21, (2), 

Spaul, E. A. 1926 A.M.N.H., (9), 17, 585-591 
Thapar, G. S- 1925 Jour. Helm., 3, 83-150 


By A. Jefferis Turner 


Mr A. J. T. Janse is at present engaged on a Revision of the whole of the world genera of this large 
family, and I am much indebted to him for sending me advance copies of some of his work. This 
has encouraged and helped me in the preparation of this paper. The family is naturally divisible into 
two subfamilies, the Anerastrianae, which have no tongue, and the Phycitinae, in which the tongue 
is fully developed. 




By A. jErrERis Turnek 
Communicated fry H. \\ J omerslev. 14 November 1946 

Mr. A. J. T. Janse is at present engaged on a Revision of the whole of the 
world genera of this large family, and I am much indebted to him for sending 
me advance copies of some of his work. This has encouraged and helped me in 
the preparation of this paper. 

The family is naturally divisible into two subfamilies, the Anerastrianae. 
which have no tongue> and the Phycilinac, in which the tongue is fully developed. 

Khy to Gexera 

1. Hindwings with 3 and 4 absent. 
Hindwings with not both absent. 

2. Forcwings with 5 absent, 3 and 4 stalked, 
Forewings with 5 absent, 3 and 4 separate, 

3. Hindwings with 5 absent. 
Hindwings with 5 present. 

4. Hindwings with 6 and 7 stalked, 
Hindwings with 6 and 7 coincident. 

5. Hindwings with cell closed. 
Hindwings with cell open. 

6. Forewings with 5 absent. 
Fore wings with 5 present. 

7. Face with conical anterior projection. 
Face not projecting. 

8. Falsi with terminal joint turned downwards. 
Palpi not so. 

9. Forewings w : th 4 and 5 stalked. 
Forewings with 4 and 5 not stalked. 

10. Kin-'wirgs wirti cell less than one-h::U. 

Hindwings with cell more than one-half. 


A I! oca 






Gen. Staitina Rag. 
N Amer. Phycit., 19. 

Face with anterior tuft of scale?. Pa'pi long, down-curved. Maxillary palpi 
minute. Forewings with 3 and 4 stalked. 5 absent. Hindwings with 3 and 4 
absent, 6 and 7 stalked. Type, S. roseolmctdla, from North America. 

Staitina rhodokaphella Rag\ (n 
Nov. Gen., 50; Rom. Mem., 8, 417, 

Queensland. New Guinea. Celebes. 

t 1 ) =z Sp. unknown to me. 
Trans. Roy. Sue. S. Aust., 71 CD, 25 July 1947 


Gen. Olamotrofha Hmps. 
P.Z.S., 1918,91. 

Face with pointed conical prominence. Palpi long, down-curved. Maxillary 
palpi dilated. Forewings with 3 and 4 separate, 5 absent, lliudwmgs with 
3 and 4 absent, 6 and 7 stalked. Type, C. pulvcrulcnta limps. 

Calamotropha ju;lverivexa limps/ *> 
P.Z.S., 1918, 91. 

West Australia: Sherlock River. 

Gen. Akerastrja Hb. 
Verz., 367. 

Face with conical anterior projection, Palpi long. ponect. Antennae of male 
with sub-basal dorsal noich. Fure wings with 5 absent. 9 and 10 stalked or 
separate. ilindwings with cell shorl, 3 and 4 stalked, 5 absent. Type, 
A. lotcih Fib., from Europe. 

Anerastria :-.iiraiul£lla Meyr. 

Proc. Linn. Soc. N.S.W.., 1878, 213. 


Anerastria erasmia Turn. 
Proc. Roy. Soc. QId. ( 1912, 117. 

Atherton, Tnjune, Carnarvon Range. 

Anerastria virginella Meyr. 

Proc. Linn. Soc, N.SAV., 1880, 233; ncurka Turn. Proc Rov. Soc. Old., 1912, 
Darwin, Brocks Creek, Cape York, Cairns, Atherton, Duaringa, Lismorc. 

Anerastria rhodochros n. sp, 
pohaxpais, rosy. 

$ ?. 18-22 mm. Head and thorax dark reddish or rosy-grey. Palpi 6; 
rosy-grey. Antennae pale grey; in male with a sub-basal dorsal notch followed 
by a fusiform glandular swelling. Abdomen grey-wlriish. base of dorsum 
ochreous. Fore wings with costa rather strongTy arched, apex rounded; bright 
rosy with slender white lines on veins confluent towards base; cilia white, partly 
rosy-tinged. Ilindwings grey-whitish; cilia white. Fore, wings broader than in 
A. virginctla Meyr,, wilh apices more broadly rounded. 

North Queensland: Cape York in September and April (W. B. Barnard); 
three specimens. Type in Queensland Museum. 

Anerastria albivena n sp. 

alhivcints, white-veined. 

$ . 19-20 mm. Flead and thorax dull rosy ; face with strong conical 
anterior projection. Palpi 6; dull rosy. Antennae pa!e grey; in male with basal 
glandular thickening. Abdomen grey, base of dorsum ochreous; tuft whitish. 
Forewings with ensta gently arched, apex rounded; bright rosy; veins slenderly 
and discretely outlined with white; cilia rosy. Ilindwings and cilia grey. 

New South Wales: Murrurundi in October (Dr. B. L. Middleton) ; one 

P) = Sp. unknown lomc 


Proc. Linn. Soc. N.S.W., 1923 T 453. 

Ankrastria .METALLAcTfs Mcyr. 

Mcyr. Tram. Ent. Soc, 1887, 262. 

Darwin, Noosa s Chinchilla, Bathurst. 

Anerastria xylodes n. sp. 

^vAwS^s-, wooden* 

3 . 24 mm. Head and thorax fuscous. PaTpi 6; pale brownish. Antennae 
grey. Abdomen oehrcous; terminal segments and tuft grey-whitish. Forewings 
elongate, costa gently arched,, apex rounded, termen obliquely rounded; pale 
brownish; a subcostal whitish stripe from base to two-thirds, edged beneath by a 
fuscous line ; a terminal scries of minute dark fuscous dots ; cilia pale grey. Hind- 
wings grey ; cilia whitish. 

North Queensland: Cairns in December (F. H. Taylor) ; one specimen, 

Anerastria clepsiphronica n. s;>. 
tcXcfrtppfiyuc'oX) deceitful. 

$, 18 mm. Head and thorax rosy. Palpi 4; rosy. Antennae grey. 
Abdomen pale ochreous. terminal segments and tuft whitish. Forewings with 
costa gently arched, apex rounded, termen obliquely rounded; 3 and 4 closely 
approximated at origin; rosy; all veins slenderly outlined with whitish ; a sub- 
marginal series of minute fuscous dots; cilia pale rosy. Hindwings and cilia 
whitish. A close mimic of Lioprosopa thlorogramma Meyr., but readily dis- 
tinguished by its shark-like head. There is a slight variation in the forewing 
from that usual in Anerastria. 

Queensland : Brisbane in October ; one specimen. 

Gen. Lioprosopa now 
AewTrpwtfOTTfK, smooth-faced. 

Face smooth or with an anterior superior tuft. Forewings with 5 absent. 
Hindwings with cell less than one-half, 3 and 4 stalked, 5 absent. Type, 
L, chlorogramma Meyr. 


Proc. Roy. Soc. Qld., 1912, 111. 

Darwin, Melville Island, Brocks Creek. 

Lioprosopa Turn. 
Proc. Rov. Soc. Qld., 1903. 117; plcurochorda Turn. Proc. Roy. Soc. Qld., 1912, 

Yeppoon, Duarmga, Brisbane, Stanthorpc. 

Lioprosopa nipiioskma Turn. 
Proc. Roy. Soc. Qld., 1912, 112. 

Darwin, Adelaide River. 

Lioprosopa kolopiiaea Turn. 
Proc. Roy. Soc. Old., 1923, 42. 
Darwin, Brisbane. 


Ltotrosopa icasmopis Turn. 
Proc. Roy. Soc. Qld. f 1903, 116. 


Proc. Roy. Soc. QU., 1904, 41. 

Thursday Island, Dalby, Stanthorpc, 

Lioprosopa eurvsttcha Turn. 

Proc. Roy Soc. Qld., 1903, 119. 
Brocks Creek, Townsville. 

Lioprosopa dimochla n. sp. 
iifi^Aot, two-barred. 

3 . 20 mm. Head and thorax greyish-brown, (Palpi missing-.) Antennae 
m male with sub-basal notch followed by a fusiform glandular swelling. Abdomen 
whitish-oehreous. Forewings with costa straight, apex rounded; brownish-grey; 
costal edge fuscous ; a whitish costal stripe from base to apex ; a similar dorsal 
stripe, from base to tornus, intermixed by fuscous dots at one-third and two- 
thirds; cilia grey, bases whitish. Hind wings and cilia whitish. 

North Australia: Brocks Creek in January (T. G. Campbell) ; one specimen. 

Lioprosopa phaulodes n. sp. 

<£tu>A">%s-, paltry. 

£ . 16 mm. Head and thorax grey. Palpi 2 and a half; grey. Antennae 
pale grey; in male with a sub-basal dorsal notch followed by a fusiform glandular 
swelling Abdomen whitish-grey, base of dorsum ochreous. Forewings with 
costa slightly arched, apex rounded; grey-whitish with sparsely scattered fuscous 
scales; a terminal series of blackish dots; ciba grey-whitish. Hindwings and 
cilia whitish. 

North Queensland: Dunk Island in May; one specimen. 

Lioprosopa pelopa n. sp. 

TreXt/jros, dusky. 

9. 18 mm. Head and thorax dark brown. Palpi 4; dark brown, base 
beneath white {Antennae missing.) Abdomen fuscous, base of dorsum brownish; 
tuft white. Forewings with costa slightly arched, apex rounded ; brown; dorsum 
broadly fuscous; costal edge white; two slender outwardly oblique white lines 
from cogla before apex; a blackish terminal line; cilia whrre, apices except on 
tornus fuscous, a short blackish median line at apex. Hindwings pale grey; cilia 

North Australia: Darwin in December (G. F. Hill) ; two specimens. 

Lioprosopa phaeochiton n. sp. 

<baioxtTwv t dark-robed. 

2. IS mm. Head and thorax fuscous. (Palpi missing.) Antennae grcy- 
whiliph. Abdomen pale grey. Forewings with costa straight, apex rounded; 
dark fuscous; a broad whitish costal stripe from base to apex, narrower towards 
base, containing very fine louditudinal fuscous lines; a tcrmmal series of minute- 
blackish dots; cilia grey with fuscous antemediau Ime. Hindwings and cilia 

North Queensland: Cape York in October (W, B. Barnard) ; one specimen. 


Lioprosopa pachyzancla n. sp. 
7ra^|ay«:Aos\ with thick sickles. 

9. 16-18 mm. Head and thorax greyish-brown. Palpi ascending, recurved, 
thickened with appressed scales, apex pointed; dark fuscous. Antennae grey. 
Fore wings narrow, cosfa almost straight, apex rounded; grey; a broad white 
dorsal streak, narrowing towards base and apex, containing very fine fuscous 
longitudinal lines and a dot at two-thirds; cilia grey. Hind wings aud cilia grey. 

North Queensland: Cape York in October and November (W. H. Barnard) ; 
five specimens. Type in Queensland Museum. 

LioPKOiQPA kyssi^ja Turn. 
Pmc. Roy. Soc. QU,. 1912. 113. 
Darwin, Chinchilla, Kmtberley. 


Proc. Roy. Soc. QTd., 190,1 117. 

Darwin, Brocks Creek, Cktudie River, Townsville, Brisbane. 

Lioprosopa sporadic?, n. sp. 
vTropaGMos, sprinkled. 

6 , 9 . 16-20 mm. TTead and thorax whitish. Palpi 3; whitish. Antennae 
pale gr?y. Abdomen whitish, base of dorsum ochreons-tinged. Forewings with 
costa straight to middle, thence strongly arched; whitish lightly sprinkled with 
fuscous and brownish ; sometmes a dark fuscous dot in disc at two-thirds ; a 
terminal series of dark fuscous dots; cilia grey, llindwinep and cilia white. 
Himlwings grev-whitish J cilia whitish. 

North Queensland: Cape. York In October and November (W. B. Laniard); 
two specimens. 

Licprcsopa rhadinodes n, sp. 
/itt-W^s, slender, 

$. 9 mm. Head and thorax white. Palpi 3; white. Antennae whitish ; 
in male with sub-basal dorsal notch. ( Abdomen missing.) Forewings with co^ta 
skglrdv arched, apex rounded; white; cilia white. 1 find wings and cilia white. 

North Queensland: Dtf;& Tdand in May; one specimen, 

Lioprosopa tarvybela n. sp. 
rai^/V*;, with long palpi. 

£ , 9 . 14-17 mm. Head and thorax grey. Palpi in male 5. in female 8; 
grev. Abdomen pale grey, hasp of dorsum ochreous. Forewings slightly arched, 
ape^:. rounded; whitish or grey-whitish ; sometimes a few subdorsal or subterminal 
blackish tkoHl z terminal series of blackish dots; cilia coneo!orous\ Hind wings 
and cilia white. 

North Queensland: Liudemaii Island in September; four specimens. 

Lioprosopa haploa n. sp. 
mttAoos. simple 

6 - 1S-20. Head giey ;. face in male whit*, in female grey. Palpi in male 3. 
white; in female 8, grey. Antennae grey ; in male dentate, with a s;uh-basal dorsal 
twitch followed by a glandular swelling. Abdomen ochreous; tuft white. Fore- 
wings wilh casta gently arched, apex rounded; grey-whitish; veins outlined wiidi 
white; soiriet'ines a %;vey line above middle from base to apex; cilia wline, liind- 
wings pale grey; cilia whits. 

North Queensland: Lindeman Island in September; four specimens. 


Lioprosopa maucida Turn. 

Proc. Linn. Soc. N.S.W., 1923, 455. 
Queensland : Miles. 

Lioptu)sopa mAcroju*uyxc;1a Turn. 
Troc. Linn. Soc N.SAV., 1923, 456. 
Townsville, Mihuerran. 

Lioprosopa thiomochla n. sp. 
fetaj/i4jg(jftj sulphur-barred. 

c t 9. 20-22 mm. Head and thorax grey. Palpi I ; grey, Antennae in 
male shortly pectinate (1 and a half); grey-whitish. Abdomen whitish-nehreous, 
base qrl dorsum brown. Forewings with costa slightly a-ehed, apex rounded ; 
brownish-grey; dorsum suffused with whitish-ochreous ; a pale yellowish crystal 
stripe from base lo apex; a fuscous dot above two-fifths dorsum; a subterminal 
line of minute fuscous dots; cilia grey. Ilindwings and cilia grey-whitish. 

North Australia: Darwin in November. Queensland; Dalby. North-West 
Australia; Kimberlcy in February, Three specimens. 

Lioprosopa colobela n. sp- 
xoAo/feAu?, with short paTpL 

<2 . 23 mm. Head and thorax pale reddish. Palpi short (I and a fourth), 
broad an base, gradually attenuating to an acute apex ; reddish-grey. Antennae 
pale, grey; In male shortly laminate, with a shallow posterior excavation clothed 
with long hairs near base. Abdomen grey; tuft ochrcotrs-whitish. Forewings 
with costa almost straight, apex obtusely pointed; dull rosy, paler towards 
tennen; veins slenderly outlined with white; a white costal stripe narrow at base 
and apex; cilia whitish. Hindwings and cilia whitish. 

North Queensland: Gordotwalc, near Glints; one specimen. 

Lioprosopa platymochla n. sp. 
TrXarvftoxXu^ broadly striped. 

3 , 25 mm. Plead and thorax white. Palpi 8; external surface grey; internal 
white. Antennae fuscous; in male shortly pectinate. Abdomen ochrcous ; tuft 
white. Forewings with costa straight, apex round-pointed; grey finely sprnkled 
with white; a broad white costal stripe from base nearly to apex; costal edge 
grey-sprinkled; a grey line beneath costal stripe; cilia white; Hindwdngs and 
cilia white. 

Queensland: Cunnamulla in October; one specimen, 

Lioprosopa transecta n. sp. 
transectus, cut across. 

22 mm. Head, antennae, and thorax white. (Palpi missing.) Abdomen 
ochreous-grey, towards base ochrcotts-brown. Forewings elongate, costa slightly 
arched, apex rounded, termen obliquely rounded; white; a dark fuscous sub- 
costal stripe from base to apex; a subterrninal series of small dark fuscous marks; 
a terminal series of minute terminal dots; cilia white. Hindwings and cilia white. 

Queensland; Tnjune in February (W. B. Barnard); one specimen. 

Lioprosopa marcioa Turn. 

Proc. Lmn. Soc. N.S.W., 1923, 455. 


Proc. Linn. Soc. N.SAV., 1923, 456. 
Townsville, Milmerran. 


Rom. Mem., 8. 397. Turn.,, Proc. Linn. Soc. N.S.W., 1923, 455. 

Darwin, Cooktown, Duaringa, Jericho, Brisbane, Sherlock River, Kimberley. 


Proc. Roy Soc, Qld., 1912, 112; arroplwea Turn., Proc. Roy. Soc. Old., 1912, 117- 
Darwin, Dunk Island, Wyndham, Timor Laut. 

Lioprosopa euryzona Mcyr. 
Etit. Mo. Mag., 19, 256. 

South Australia ; Wirrabara, 

Lioprosopa acidnias Turn. 

Proc. Roy, Soc, 1903, 117. 


Lioprosopa minimella Hmps. 

Rom. Mem., 8, 392; Turn., Proc. Linn Soc. N.S.W., 1923, 454. 
Darwin, Thursday Island, Celebes, Borneo. 

Lioprosopa chlorocramma Meyr. 

Proc. Linn. Soc. N.S.W., 1899, 1,116; rhodosticha Turn., Proc. Linn. Soc. Roy. 
Soc. Qld., 1903, 116. 
Reid River, Duaringa, Brisbane, Tweed Heads, Maepherson Range, Rose- 
wood, Toowoomba, Injune, Milmerran, Murrurundi. 

Lioprosopa poliosticha n. sp. 
TruXioirrixo^ grey-lined. 

6 , 22-24 mm. H^ad and thorax whitish. Palpi 4 and a half; grey-whitish. 
Antennae whitish; in male with a sub-basal dorsal notch. Adhomen whitish, basal 
half of dorsum ochreous; tuft wlute. Forewings with costa gently arched, apex 
obtuse ; white lightly sprinkled with minute grey scales ; a pale grey median stripe 
from base beneath cell to vein 2; cilia white. Bindwings pale grey; cilia white. 

North Queensland: Herherton in September; four specimens. 

Lioprosopa rhantista n. sp. 
avmros-, sprinkled. 

i . 24-26 mm. Head and thorax grey. Palpi 3; whitish. Antennae whitish- 
in male with a sub-basal comb of long hairs. Abdomen ochreous or fuscous ; tuft 
white. Forewings with costa rather strongly arched, apex rounded; white very 
lightly sprinkled with pale grey except on veins; cilia white. Hindwings grey- 
whitish ; cilia white. 

North Queensland: Lindeman Island in September; two specimens. 

Lioprosopa t.arqpis Turn. 

Proc. Roy. Soc. Qld., 1912, 113. 

Lioprosopa argosticiia Turn. 

Proc. Roy. Soc. Qld., 1912, 115. 
Darwin, Cape York, Dalby. 


Lioprosopa fsamatiirlla Meyr. 

Proc. Lion. Soc. N.S.W., 1879, 234; nilcus Ruth, Tr. Eni. Soc, 1886, 440; baliora 
Turn., Proc. Roy. Soc. Old., 1912, 116. 

Cairns, Peak Downs, Brisbane, Dalby, Sydney. Fernshaw. 


Proc. Roy. Soc. Qld., 1912, 116. 


Lioprosopa abixpta Turn. 
Proc. Roy. Soc. Old.. 1912, 114. 

Darwin. Cairns, Dnnk Island, Reid River, Nambour, Brisbane, Mount Tam- 
borine, Bunya Mountains. 

Lioprosopa taliella limps. 
Rom. Mem., 8, 402. 

Lioprosopa mixorat.ts Low. 
Trans. Roy, Soc. S. Aust., 1903, 52. 

Gen. Anchylobela nov. 
n-/xv^o/3e\o^ t witb crooked palpi. 

Tongue absent. Palpi moderate, porrect; terminal joint bent downwards. 
Antennae in male with sub-basal dorsal notch, followed by a fusiform swelling. 
Forewings with cell two-thirds, 2 from before angle. 3 and 4 stalked, 5 absent. 
Hindwings with cell one-half, 2 from before angle, 3 and 4 stalked, 5 absent. 

Anchylobela haplodes n. sp. 

uttA&A^, simple. 

$ . 16-17 mm. Head and thorax pale ochreous grey, Palpi 2 and a half, 
terminal joint short, pointed; pale ochreous-grey. Abdomen pale ochreons-grcy, 
towards base deep ochreous. Forewings with rosta arched,, apex rounded, termen 
obliquely rounded; pale ochreous-gre-y with very scanty fuscous sprinkling; a 
terminal scries of blackish dots; cilia whitish. Hindwings grey-whitish ; cilia 

North Queensland : Kuranda in SepLember (W. B. Barnard), two specimens. 
Type in Queensland Museum. 

Gen. Saluiua Rag. 

Palpi long, porrect. Antennae in male with sub-basal dorsal notch. Fore- 
wings with 4 and 5 stalked. Hindwings with cell less than one-half. 3 and 4 
stalked, 5 absent. 

Salurta aoknocera Turn. 

Proc. Linn. Soc. N.S.W., 1923, 458. 
Cairns, Cunnamulla. 

Saluria rhopoessa Turn. 

Proc. Roy. Soc. Qld., 1903, 120; distichdla limps., Proc. Zool. Soc, 1918, 101. 
Brocks Creek, Cairns, Townsville. 


Proc. Roy. Soc. Old., 1903, 120, 

Claudie River, Palm Island, Townsville. 


Saluria leucoxeura Turn. 

Proc Roy. Soc Qld., 1912, 118. 

Darwin, Cape York, Innisfail, Townsviilc, Ayr, 

Saluria iiqlochroa Turn. 

rroc. Roy. Soc. N.S.W., 1903, 121. 

Duaringa, Emerald, Birchrp. 

Saluria pleurosticha Turn. 
Proc, Roy. Soc. Qld., 1903, 115. 
Brocks Creek, Townsville. 

Saluria ctenucha 

Proc. Roy. Soc. Old., 1912, 118. 
Darwin, Townsville, Injun e. 

SalukiA grammivkna Pimps. 
Proc. Zool. Soc, 1918, 99; Turn., Proc. Linn. Soc. N.S.W., 1923, 459. 
North Australia: Alexandria, Sherlock River. 

Saluria stereochorda n. sp. 

(TTepeoxophos, straight-barred. 

t! , 9 . 24-26 mm. Head white. Palpi in male 3 and a half, in female 5; 
grey, lower edge white. Maxillary palpi in male short, ending in a terminal tuft 
of long white hairs; in female filiform; grey. Antennae grey. Thorax white; 
bases of tcgulae grey. Abdomen in male grey; in female ochreous; in both with 
base and tuft white. Forewings with costa straight, apex rounded; white, in 
female sprinkled with grey; a straight subcostal grey bar from base to apex in 
male, m female a median suffused bar extending to angle of cell ; veins 2 to 5 
grey; in male cilia white, but on apex grey; in female grey with white apices. 
Hindwings grey; cilia white. 

Queensland: Bunya Mountains in September; two specimens. 

Saluria pelochroa n. sp. 

7reAo'x/>otw. dusky. 

9 . 2S mm. Head, thorax, abdomen, and antennae fuscous. Palpi 3, 
ascending; fuscous. Forewings with costa straight, apex rounded; fuscous; a 
whitish line sprinkled with fuscous on costa from base to apex, 
Tasmania: Burnie in January; one specimen. 

Saluria desertella Hmps. 
Proc. Zoo!. Soc, 1918, 977; Turn., Proc. Linn. Soc. N.S.W., 1923, 459. 

Alexandria, Injune, Carnarvon Range, Charlcville, Cunnamulla, Sherlock 
River, Wyndham. 

Saluria xiphomela Low. 

Trans. Roy. Soc. S. Aust., 1903, 52. 

Saluria neotqmella Meyr, 

Proc. Linn. Soc. N.S.W., 1879, 226. 


Saluria neuricella Hmps. 
Proc. Zool. Soc, 1918, 98. 

Peak Downs. 

Saluria leuconeukklla Hmps. 
Rom. Mem., 8, 339. 

Gen. Emmalocera Rag. 

Nouv. Gen., 38; Turn., Proc. Linn. Soc. N.S.W., 1923, 460, Type, U. leucocincta 
Wlk., from Archipelago. 

Emmalocera lokgiramella limps. 
Rom. Mem., 8, p. 460. 

Darwin, Brocks Creek, Towtisville, Ayr, Brisbane, 


Bull. Soc. Eat Fr., 1869, 220; rodiaiclla Hmps.. Rom. Mem., 8, 315; rhabdoid 
Turn., Proc. Roy Soc. Qld.. 1903, 122; achrosia Turn., Proc. Roy. Soc. 
Qld., 1903, 122/ 
Cape York to Tweed Heads, Buuya Moun.ains, Stanthorpc, Tabulum, Ben 
Lomond, New Guinea. 

Emmalocera achromatella Hmps. 

Proc. Zool. Soc, 1918, 126. 

Dalby, Charleville, Broken Hill. 

Emmalocera crossospila tu top* 
KpouvofTTTiXos, with marginal dots. 

5. 24 mm. Head and thorax pale ochreous-grey. Palpi 8; pale grey. 
Antennae grey.. Abdomen grey-whitish, partly ochreous-tinged ; twit whitish. 
Forewings with costa gently ached, apex acute, mermen straight, oblique; whitish- 
ochrcous; a blackish discal dot above middle at three-fifths; a terminal series^ of 
blackish dots; cilia white with a grey median line. Hindwings and cilia white. 

Queensland: Yelarbon in November; one specimen. 

Gen. Anarcsca nov. 
■paptiSKo-;, unattractive. 

Palpi slender, porrect. Forewings with 2 from well before angle, 3 from 
angle, 4 and 5 approximated at origim 8 and 9 stalked, 10 from near end of cell. 
Hindwings whh'cell open, 3, 4, 5, stalked out of 2, 7 anastomosing with 12. 

Anaresca xuthochroa n, sp. 
£ov0oxpt»"ov. yellowish , 

2 . 24 mm. Head and thorax whitish-ochreous. Palpi 2; grey. Antennae 
grey. Abdomen ochreous ; tuft whitish. Forewings wi:h costa straight, apex 
rounded-rectangular ; ochreous ; a whitish median line from base _ expanded 
towards tcrmen; a terminal series of blackish dots; cilia whitish with a grey 
median line. 

North Queensland: Lindcmau Island in September; one specimen. 

Gen. Alloea nov. 
rlAAnMK, different. 

Labial palpi short, obliquely ascending. Maxillary palpi about half length 
of labial. Face with acute conical projection. Forewings with 2 from before 
angle, 3 from angle, 4 and 5 stalked, 8 and 9 connate or short-staked, 10 from 
near end of cell. PImdwmgs with 5 absent. 3 and 4 approximated from angle of 
cell, 6 and 7 coincident and anastomosing with 12; cell two-thirds. 


Alloea xylochroa n. sp. 
fi'Aityaoos, wood-coloured, 

$. 26-28 mm, Head and thorax pale brownish-ochreous. Palpi 2; pale 
brownish-ochreous, lower edge white. Antennae grey. Abdomen pale ochreous ; 
extreme basal and apical segments white ; tuft ochreous. Forewings with costa 
slightly arched, apex rounded; pale ochreous with fuscous markings; sometimes 
an interrupted sub-basal transverse line; sometimes an interrupted posi median 
blackish line; a white dot beneath two-thirds costa; a dentate subterminal line, 
sometimes interrupted, sometimes broadly suffused anteriorly; followed by a 
dentate whitish line; a triangular subapical fuscous spot narrowly prolonged lo 
near fornus; a terminal series of dark fuscous dots; cilia whitish with a median 
fuscous line, Hindwings grey-whitish; cilia whitish. 

North- West Australia: Wyndham in January (T. G, Campbell) ; two 

Gen. Creocota Turn. 
Froc. Linn. Soe. N.S.W., 1931, 342. 

Tongue absent. Labial paipi obliquely ascending, slightly curved upwards; 
second joint with appresscd scales, rather slender; terminal joint short, conical, 
obtusely pointed. Maxillary palpi large, in male ending in a tuft of long hairs. 
Forewings with cell two-thirds, discocellulars incurved. 2 from angle well 
separated from 3, 3 much nearer 4 than 2 at origin, 4 and 5 stalked. Hindwings 
with cell one-half, discocellulars incurved, the lower very oblique, 2 from three- 
fourths, 3 from angle, connate with 4, 5, which are stalked. Monotypical. 


Proc. Linn. Soc. N.S.W., 1931, 343. 

Key to Gexrra 

1. Himiwincs with 4 and 5 absent. 
Hindwings with 4 and 5 not both absent. 

2. F'orewings with 8 and 9 coincident, hhicUvings with cell open. 
Forewings with 8 and ° stalked, liimhvin^s with cell dosed. 

3. Forewings with 3 and 4 stalked. 
Forewings With 3 and 4 connate. 

4. Hindwings with 4 present, 5 absent. 
Hindwings with 4 and 5 present. 

5. Forewings with 9 absent. 
Forewings with 8 and 9 stalked. 

6. Forewings with 5 absent. 
Forewings with 5 present. 

7. Palpi ascending, 
Palpi porrect. 

8. Forewings with 8 and 10 stalked, 
Forewings with 8 and 10 separate. 

9. Palpi ascend iojj. 

Falpi porrect. 

10. Forewings with 5 absent. 
Forewings with 5 present. 

IF Forewings with 3 and 4 stalked. 
Forewings with 3 and 4 separate. 

12. Hindwings with cell one-fifth. 

Hindwings with cell one-third to one-half. 





























13. Palpi ascending, recurved. 
Palpi straight. 

14. Hindwings with cell one-half. 
Hindwings with cell open. 

15. Palpi swollen to enclose penicillate maxillary palpi. 
Palpi nut swollen, maxillary palpi filiform. 

16. Hindwings with 3 and 4 connate or stalked. 
Hindwings with 3 and 4 separate. 

17. Forewings with 2 and 3 stalked. 
Forewings with 2 and 3 not stalked. 

18. Hindwings with 2 from "well before angle of cell, which is long. 
Hindwings with 2 from or from near angle of cell, which is abort, 

19. Forewings with 4 and 5 stalked. 
Forewings with 4 and 5 separate. 

20. Palpi ascending. 
Palpi porrect. 

21. Forewings with 4 and 5 stalked. 
Forewings with 4 and 5 separate. 

22. Forewings with 5 from above angle of cell well separated from 4. 
Forewings with 4 and 5 approximated at origin. 

24. Palpi ascending. 
Palpi porrect. 

25. Maxillary palpi filiform. 
Maxillary palpi in male penicillate. 

Gen. Ernophthora Mcyr. 
Trans. EnL. Soc, 1887, 263. 

Palpi ascending, recurved. Forewings with cell from near angle, cell open, 
3 and 4 stalked, 5 absent, 8, 9, 10 stalked. Hindwings with cell open, 4 and 5 
absent. Type, E. phoenicias Meyr. 

Ernophthora phoenicias Meyr. 

Trans. Ent. Soc, 1887, 263. 


Erkophthora milicha Turn. 

Proc. Unn, Soc, N.S.W., 1931 , 342. 
Yeppoon, Macphcrson Range. 

Gen. Dialkpta Turn. 

Proc. Roy. Soc. Qld., 1912, 119. 

Palpi ascending, recurved. Forewings with $ and 4 stalked, 5 absent, 8 and 9 
coincident. Hindwings with cell closed, 4 and 5 absent. Type, D. micrapolia 

Dialepta micropolia Turn. 

Proc. Roy. Soc. Qld., 1912, 119. 
Cairns, Brisbane. 

Gen. Abarys now 

afjapvi, light. 

Falpi ascending, recurved. Forewings with 3 and 4 connate, 5 absent, 8 and 9 
stalked. Hindwings with cell closed, 3 and 4 absent, 2 and 5 widely separate. 


Abarys amaurodes n. sp. 
apavfitaSijSt obscure. 

2 . 16 mm. Head, palpi, anlcnnaej and thorax fuscous. Forewings narrow, 
posteriorly dilated, apex round-pointed, tenncn straight, oblique; grey lightly- 
sprinkled with fuscous; marking's dark fuscous; an outwardly oblique fascia at 
two-fifths, paler in centre; a transverse subcostal mark at two-thirds; a terminal 
suffusion and cilia fuscous. Hindwings and cilia grey. 

Queensland: Brisbane in September; one specimen. 

Gen. Kphestia Gn. 
Fur. Micro,,, 81. 

Palpi ascending, recurved. Forewings with 5 absent, 9 absent. Hindwings 
with cell nearly reaching middle, 3 arid 4 closely approximated or stalked, 5 absent. 
Type E. ehitella lib. 

Epiikstia lib. 
Meyr. Rev. Hdbk. Brit. Lep., 373. 
Sydney, Gisborne, Melbourne. 

Efukstia i'xcui-ella Barrett 

Meyr. Rev. Hdbk. Brit. Lep.. 388. 

Darwin, Cairns, Gladstone, Brisbane, Dal by, Brunswick Heads, Deloraine. 
South Australia: Kadina, Africa. Europe. America. 

Ephestia pelopis n.sp. 

7rcAa)7rtti, dusky. 

5 . 19 mm. Head, palpi, and thorax greyish-brown. Antennae grey. 
(Abdomen missing.) Forewings with costa straight to middle, thence arched, 
apex round-pointed, termen slightly rounded, moderately oblique; greyish-brown ; 
a faint suffused transverse line at two-fifths; a similar but narrower line from 
beneath two-thirds costa oblique to mid-dorsum ; a terminal series of minute 
fuscous dots ; cilia pale grey, Hindwings grey-whitish ; cilia grey-whitish, apices 
whitish.; cell one-fourth. 

North Queensland; Kuranda in October (F. P. Dodd) ; one specimen. 

Ephestia cautella Wlk. 
Meyr. Rev, Hdbk. Brit. Lep., 388. 

Darwin, Atherton, Lindeman Island, Townsville, Maryborough, Brisbane, 
Birchip, Kangaroo Island, Perth. 


Meyr. Rev. Hdbk. Brit. Lep.. 388. 

Brisbane, Dunkeld, Launeesron, Adelaide. Africa. Europe. America, 

Gen. Plodta Gn. 

Meyr. Rev. Hdbk. Brit. Lep., 386. 

Palpi porrect, Forewings 2 and 3 sometimes stalk. 5 absent, 9 absent. Hind- 
wings eel! nearly reaching middle, 3 and 4 connate, 5 absent. Type, P. infer- 
puncifUa Gn. 


Meyr. Rev. Hdbk. Brit. Lep., 386. 

Mackay, Brisbane, Toowoomba, Slanthorpe, Milmerran, Cliarleville, Sydney, 
Birchip, Adelaide. Perth. 


Gen, Ecbletodes Tunj. 
Proc. Roy. See. Qld., 1903, 124. 

Palpi ascending, recurved. Fore wings with 4 and 5 stalked, 9 absent, 8 and 
10 stalked. liindwings with cell one-half, 2 from before angle. 3 and 4 stalked, 
5 absent-. Type, £. pscphcims Turn Enchryphodrs Turn.. Proc. Hoy, Soc. Qld., 
1912, 123, is a synonym. 

Ecbletodes psephexxas Tnn\ 

Proc. Roy. Soc. Old... 1903, 125. 
Brisbane, Lismore. 

Ecbletodes aknicta Turn. 
Proc. Roy. Soc. Old, 1912, 120. E. aenictopa Turn., Prcc. Roy. Soc. Old., is a 
Cairns, Atherton, Palm Island, Darwin. 

Ecbletodes OTOrnrA Turn. 

Proc. Roy. Soc. Qls., 1912,, 120. 

Gen. Homoeoso^a Curt. 

Ent. Mas., 1, 190; Hnrps. Moths JrtcL, 4, 66. 

Palpi, ascending, recurved. Eorewings with 4 and 5 staked* 9 absent. Hind- 
wings with ceil short, 3 and 4 approximated or conrate, 5 absent. Type, 
II. suwclla Fab., from Europe. 


Isis, 1848, 863. Meyr. Troc. Linn. Soe. N.S.W-, 1878. 214. 

Cairns to Victoria, Ilughcnden, Cunnamuila, Broken Hill, Birchip, Adelaide, 
Western Australia. 

Homoeoscma fornacella Meyr, 

Proc. Linn. Soc. N.S.W., 1S80, 219. 

Cairns, Macpherson Range, Ben Lomond, Ebor, Sydney, Tasmania. 

Homoeasorna centrcsticha n. sp. 

KtvTpovTixp** with central line. 

o, 9. 18-20 mm. Head and thorax grey. Palpi and antennae fuscous. 
Abdomen grey; tuft whitish. Forewings narrow, costa straight, apex pointed; 
grey; markings dark fuscous; a line from base to end of cell, connected by an 
inwardly oblique line witb one-third costa; a white subcostal line from base to 
end of cell; two fuscous dots placed transversely at end nf cell; some minute 
terminal dots; cilia whitish. Hindwings grey-whitish; cilia whitish. 

Queensland: Brisbane in October; Hunya Moumaius in February. New 
South Wales: Murrurundi in March. Three specimens. 

Homoeosoma rhapta n. sp. 
/'airro?, embroidered. 

? . 18-20 mm. Head grey; face white. Palpi grey; second joint with post- 
median and terminal fuscous bars. Antennae pale grey. Abdomen grey, towards 
base dark fuscous. Forewings with costa nearly straight., apex obtuse; grey with 
some fuscous sprinkling; markings dark fuscous; a triangular spot on base of 
costa; a narrow transverse fascia at two-fifths, becoming broader towards termen; 
a minute subcostal dot at two-thirds; a subtenninal line obtusely angled in middle 


and again between this and dorsum, edged posteriorly whitish; a terminal scries 
of dots; cilia grey, Hindwings pale grey; cilia whitish with sub-basal grey line. 
Queensland: Aramac in June; Stanthorpe in November from larva on 
Acacia (H, Jarvis). 

Homoeosoma euryleuca n. sp. 
pvXtvKo;, broadly white, 

f , 17 mm. Head fuscous-brown. Palpi exceeding vertex; brown. 
Antennae grey. Abdomen whitish -ochreous; tuft whitish. Forewings with costa 
slightly arched, apex pointed; bxownisli-grey ; a white costal stripe from base, 
gradua'Jy increasing in breadth to middle, thence broad to ternien ; two minute 
fuscou- dots placed transversely at two-thirds; cilia white. Hindwings whitish- 
grey; cilia white. 

North Queensland: Cape York in October (W. B. Barnard) ; one specimen, 

Homoeosoma contracts n. sp. 

contractus, narrow. 

?. 17 mm. Head and thorax grey. Palpi fuscous., upper edge whitish. 
Antennae fuscous. Abdomen dark fuscous v towards apex grey; tuft ocbreous- 
whitish. Forewings narrow, slightly dilated posteriorly, costa slightly arched, 
apex obtuse, tcrmen straight, oblique; fuscous sprinkled with white; a narrow 
-white line above middle from one -fifth to three-fifths ; a dark fuscous median 
line through cell ; suffused fuscous dots above dorsum at one-sixth and one-third; 
a slender oblique subterminal line parallel to termen; citia gxzy. Hindwings pale 
grey; cilia white with faint grey sub-basal line. 

New South Wale,s: Murrurundi in February (Dr. R. L. Middleton) ; one 

Homoeosoma pclosticta n. sp. 
Tre\ct<7Tu<To> t dusky-spotted. 

3 , ? . 12-16 mm. Head, thorax, and abdomen grey. Palpi and antennae 
fuscous. Forewings narrow, costa almost straight, apex obtuse; grey with a few 
minute fuscous dots ; one median and sub-basa] ; another, sometimes double, at 
end of cell; se\-eral in subterminal area; cilia white. Hindwings. grey-whitish; 
cilia white. 

Queensland: Biloela (foodplaut Sortjltinv), in October. New South Wales: 
Ehor in December. Three specimens. 

Homoecs-o-ma ochropasta n. sp. 

w^jo-air™*, pale-sprinkled. 

?. 16 mm. Head, antennae, thorax, and abdomen grey-whitish. Palpi 
grey, extreme base and apex white. Forcvings narrow, costa nearly straight, 
apex obtuse; whitish finely sprinkled with pale grey; a terminal scries of minute 
terminal dots; cilia grey-whitish. Hindwings and cilia white. 

North Queensland: Kuranda, in May ( F. P. Dodd); one specimen. 

Homoecson'a atcchna m sp. 
a-n-xrus, simple, 

9 . 21 mm. Head, thorax, and abdomen £rey. Valpi and antennae fuscous. 
Forewings with costa slightly arched, apex rounded; grey-whitish; cilia grey- 
whitish. Hindwings and ciiia grey-Avhltish. 

Tasmania: Dcrwcnt Bridge,, in January; two specimens. 


Tree. Koy. Soc. QUI., 1903, 126. 



Trans. Roy. Soc. S. Aust., 1903, 58. 

Cairns, Atherron, Brisbane, Mount Tamborine, Toowoomba, Carnarvon 
Range, Sy<lney. 

Homoeosoma lechriosema n. sp. 

A€ / \ptocrw/.f,f>s, obliquely marked. 

g . 24 mm. Head, palpi, and thorax grey. (Antennae missing.) Abdomen 
pale ochreous-grey, towards apex whitish. Forewings dilated posteriorly,, costa 
straight, apex obtusely pointed; fuscous densely sprinkled with white; a basal 
fuscous suffusion edged posteriorly by a slender oblique line from costa near 
base to one-third dorsum, and itself edged porteriorly by a fuscous line; a fuscous 
subcostal dot at three-fifths; a slender oblique white "line from costa near apex 
to three-fourths dorsum, edged anteriorly by a fuscous line; cilia grey-whitish 
sprinkled with white. Hindwings and cilia whitish. 

Queensland; Tweed Heads, in September; one specimen. 

Homgkoso-UA faiukaria Turn, 
Proc. Roy. Soc. Qld., 1903, 126. 

Cairns, Eungella, Tweed Heads, Mount Tamborine, Macpherson Range, 
Toowoomba, Bunya Mountains.. Stanthorpe, Lismore, Ebor, Sydney. Stratum, 
New Zealand. 

Gen. Syntypica Turn. 
Proc. Linn. Soc. K.S.W-, 44. 

Palpi lon^, porrect. Forewings with 4 and 5 absent, 9 absent. ITindwings 
with 3 and 4 separate, S absent. Type S. aienrodes Turn. 


Proc. Roy. Soc. Old., 1904, 45. 

Gen. CKYJt'TAoiA Turn. 
Proc. Roy. Soc. Old-, 121. Type, C. xitihobcla Turn. 

Palpi ascending, recurved. Forewings with 3 and 4 stalked, 5 absent, 8 and 
9 stalked. Ilindwings with cell short (one-fifth ) . 5 absent". Type, C< xtiihnbcla 

Cryptadta xrTiiOBEEA Turn. 

Proc, Roy. Soc. Qld., 1912, 121. 


Gen. Euzopncnorms Hviips. 
J. Bombay Soc, 1897, 475; Rom. Mem., 8, 79. 

Paipi short, ascending, recurved. Forewings with 3 and 4 connate or 
stalkcd, 5 absent, 8 and 9 stalked. Ilindwings with cell one-third to one-half, 
2 from before angle, 3 and 4 approximated, connate, or stalked from single, 
5 absent. Type, E. albicans limps. 


J. Bombay Soc, 1897, 475. 

15-16 mm. Head and thorax, ochre.ous-whitish linked with grey. Palpi, 
antennae, and abdomen grey. Forewings; elongate-triangular, costa slightly 
arched, lermcn nearly straight, oblique; whitish lightly sprinkled with fuscous; 
markings fuscous; a white line from one-third costa to one-third dorsum edged 


fuscous posteriorly; a slightly waved wlifije subtennixial line; cilia while. Hind- 
Mings grey-whitish ; cilia white. 

For early stages see Proc. Roy. Soc. Qld., 1903, 127. 

TownsvtJle, Brisbane, Brunswick Heads, Ceylon, India. 

Euzophcrodes concinella n. &p, 
CMncuincUtis, neat 

9 . 16 mm. Head, palpi, thorax and abdomen whitish-ochrcoua. Antennae 
and abdomen grey. Forcwmgs narrow, costa gently arched, apex subreetangular, 
tennen straight, slightly oblique; oehreous-wlrlHsh slightly sprinkled with fuscous; 
markings dark fuscous; basal clots on cosla and dorsum almost meeting; an oval 
dot on fold at one-fourth ; three minute dots in a transverse line at one-third; a 
sttbco^l dot at three-hubs with another slightly beneath and beyond; a sub- 
terminal line; cilia ochreous-wliitish. Hindwings and cilia grey. 

Cape York in October (W. B. Barnard) ; one specimen. 

Trails. Roy. Soc. S. Aust., 1903, 57. 

EvzopiiEKonrs ^podoftila Turn. 

Proc, Roy. Soc. Old., 1912, 121. 

EuzoPtTZRoors ixptocos-ma Turn. 

Proc. Row Soc. Qld., 1903. 127; poliocroua Low., Trans. Rov. Sue. S. Aust. t 
1905, 104. 
Townsville, Mi!merran, Broken Hilk 

Euzopherodcs homophaea n. sp. 
f'/^.o^>ato«, uniformly dusky. 

2, Head and thorax fuscous. Palpi pale brownish. Abdomen grey; tuft 
grey-whitish. Forewings narrow, slightly dilated posteriorly, costa gently 
arched, apex rounded; fuscous-brown; markings obscure, fuscous; an outwardly 
curved subtermiual line of clots nor reaching costa; some terminal dots; cilia 
grey. Hindwings grey-whitish ; cilia whitish. 

North Queensland: Kuranda, in March ( F. P. Dodd) ; one specimen. 

Eyzopherodes homocapna n. sp. 
''fiQKa7rvo>s, uniformly dark. 

Q . 16 mm. Head, palpi, antennae, thorax, and abdomen fuscous, Fore- 
wings narrow, cOsta straight, apex rounded ; dark fuscous; cilia fuscous. Hind- 
wings w:lh cell open, 3 and 4 stalked to near termen; grey, cilia grey. 
Queensland: Cunnamulla, in October; one specimen. 

Euzopherodes schematica n. sp. 
u^yy/.'ur/.xri'?, well designed. 

o. 2. 15-16 mm. Head, thorax, and abdomen fuscous; face sometimes 
partly v;hite, Palpi fuscous, base and a median ring on seeond joint white. 
Antennae fuscous. Forewings nnrrow, costa straight, apex subreetangular; white 
with dark fuscous markings; an outwardly oblique basal fascia; a second fascia 
from one-fourth costa to mid-dorsuui; a dot beneath mideosta; an oblique line 
from apex to second fascia; a subterminal line followed by some suffusion; a 
terminal series of minute dots; cilia grey. Hindwings and cilia pale grev. 

North Queensland : Cape York, in November (W. B. Barnard) ; five 
specimens. Type in Queensland Museum. 


Euzopherodes phaulopa n. sp. 

</>atv\w7ro?, mean-looking. 

9. 26 mm. Head, antennae, thorax, and abdomen grey; pectus white. 
Palpi fascous. Forewings elongate, costa gently arched, apex rounded, termen 
obliquely rounded; pale grey sprinkled with fuscous; a transverse fuscous line 
at one-fourth, angled inwards beneath costa. between this and dorsum finely 
dentate ; some terminal dots ; cilia grey. Hindwings with cell one-fourth ; 
grey-whitish ; cilia whitish. 

" New South Wales: Scone, in September (H. T. Nicholas); one specimen- 
Gen. ScythrophaneiS no v. 
aKvBpcxjiairtjs, gloomy, 

Palpi ascending, recurved. Forewings with 3 and 4 separate, 5 absent, 
8 and 9 stalked. Hindwings with cell open, 5 absent. Type, U. apatdia Turn. 

Scvrnuoi'iiAJsK'S ai>atej.ia Turn. 

Proc. Roy. Soc. Old., 904, 45. 


Proc, Roy. Soc. Old., 1912. 122. 


Pror. Roy. Soc. Old., 1912, 122. 

Gen. Unadilla Hulst. 

Trans. Amer. Ent. Soc. 1890, 197. 

Palpi ascending, recurved. Maxillary palpi filiform. Forewings with 3 and 

4 separate. 5 absent, 8 and 9 .stalked. Hindwings with cell one-half. 3 and 4 
stalked, 5 absent. 

Unadtlla uIvSttchella Meyr. 

Proc. Linn. Soc. N.S.W., 1878, 215. 

Brisbane to Gisborne, Glen Innes, Tastnania, 

Gen. Ancvlodes Rag. 

Ann. Soc. Ent. Fr., 1887, 250. 

Palpi straight, ascending, exceeding vertex; second and terminal joints 
strongly dilated, apex obtuse. Maxillary palpi of male ending in a pencil of long 
hairs. Antennae in male with scape dilated, base of shaft strongly dilated 
antero-posteriorly. Forewings with 2 from before angle.. 3 and 4 from angle, 

5 absent, Hindwings with 3 &nd 4 connate, 5 absent. 

Ancylodes feniciixata Turn. 

Proc. Roy. Soc. QM., 1907, 46. 


Gen. Eucamtyla Mcyr. 

Proc, Linn. Soc. N.S.W., 1S82, 159. 

Palpi moderate, porrect. Forewings with 4 and 5 stalked, 7 and 8 stalked- 
Plindwings with 3 and 4 approximated at origm, 5 absent. Type, E. ctheicUa 


Proc. Linti. Soc. N.S.W., 1882, t7l. 


Gen. Crocyuopora Meyr. 
Proc. Linn. Soc. N.S.W., 1SS2, 158. Type, C. clnigerclla Meyr. 

Palpi long, porrect Maxillary palpi filiform, Forewings with 2 from 
before angle, 3 and 4 closely approximated from angle, 5 absent. 8 and 9 stalked. 
Hindwings with 2 from before angle, 3 and 4 connate, 5 absent; cell some- 
times open, 


35, 1719; sienoptcrella Meyr., Proc, Linn. Soc. N.S.W., 1878, 200. 

Atherton, Mackay, Duaringa to Victoria, Glen I, Charleville, Ebor, 
Broken Bill, Birchip, Strahan, Mount Lofty, Western Australia, Norfolk Island, 
New Zealand. 

Gen. Mekeintadia 

Palpi ascending", recurved. Forewings with 2 and 3 stalked, 4 and 5 stalked, 
8, 9, 10 stalked, llindwings with 2 from near angle, 3 and 4 stalked, 5 absent. 
I am unable to give the origin of this name, which I received from Sir Geo. 


30, 958. 

12-13 mm. Head, palpi, antennae, thorax, and abdomen ochreous- whitish. 
Forewings narrow, costa straight, apex obtuse, termeu obliquely rounded; 
ochreous-whitish with fuscous markings; a dot on base o[ costa; a short dorsal 
streak from base; a longer streak on fold; a slender oblique streak from two- 
fiEths costa, sometimes connected with a longitudinal streak above middle ; some- 
times another streak below middle ; both ending in a large terminal suffusion ; 
cilia fuscous. Hindwings grey-whitish ; cilia vvhitish. 

North Queensland: Cairns, Innisfail. 

Gen. EtjzoPheha Zel. 
Trans. Ent Soc, 1867, 453. 

Palpi ascending, recurved. Forewings with 4 and 5 stalked, 8 and 9 stalked. 
Hindwings with 3 and 4 stalked. 5 absent. Type, E r hivlclla ZiL, from Europe. 

Euzopiieka subakcl'ella Meyr. 
Proc. Linn. Soc. N.SAV,, 1878, 211, 

Darwin, Cape York to Victoria, Glen Innes, In June, Adavalc, Sea Lake. 
Mount Lofty. Artlrossan, Perth. 

EuzorHEFA aoropHRACMA Meyr. 

Trans. Ent. Soc, 1887, 255. 


Euzophera albiccsta n. sp. 
alhicasliis, with white costa. 

% , 9 . 18-22 mm. Head fuscous with narrow lateral margins. Palpi 
fuscous, basal half white. Antennae and thorax fuscous. Abdomen grey; apices 
of segments and tuft white. Forewings narrow, costa straight, apex rounded ; 
grey; u white costal line from to near apex; more or less broadly suffused 
and sprinkled with grey; a transverse fuscous fascia at one-third; an oblique 
fascia from apex to three-fourths dorsum, edged with white posteriorly; cilia 
grey. Hindwings and cilia pale grey. 

Western Australia: Denmark in March;, Yanchep in September; four 


Euzophera flavicosta n. sp. 
flavicostus, with yellow costa. 

9 . 18-20 mm. Head, palpi, thorax and abdomen brown. Antennae fus- 
cous. Forewings with costa straight, apex rounded, termen oliquely rounded; 
fuscous-brown; a broad yellowish costal line from to near apex, interrupted 
by a yellowish dot at three-fifths; a slender slightly dentate subtermmal line; 
terminal edge yellowish interrupted by a series of fuscous dots; cilia grey. Hind- 
wings grey-whitish; terminal edge fuscous; cilia grey-whitish with a sub- 
basal grey line. 

North Queensland: Cape York in October (W. B. Barnard); four speci- 
mens. Type in Queensland Museum, 

Euzophera ischnopa n. t,p> 
ttirVyfciintej thin. 

2 . 18 mm. Head and thorax dark fuscous. Palpi much exceeding vertex; 
dark fuscous, extreme base whitish. (Antennae missing.) Abdomen ochreous; 
tuft paler. Forewings narrow, slightly dilated posteriorly, costa gently arched, 
apex obtuse; fuscous-brown; three fuscous dots in cell and two placed trans- 
versely at its end; cilia fuscous. I find wings pale grey; cilia grey-whitish with 
a faint sub-basal grey line* 

North Queensland: Cape York in October (W. B. Barnard) : one specimen. 

Euzophera arrbythmopis n. sp. 

dppvOfmnt s% di so rde rly . 

9 . 12-14 mm. Head, palpi, antennae, and thorax grey. Abdomen fuscous; 
tuft whitish. Forewings dilated posteriorly, costa straight to three-fourths, thence 
ached, apex rounded ; grey; an outwardly curved whitish line from one-third costa 
to mid-dorsum preceded by some fuscous suffusion; a hue whitish subtcnninal 
line indented in middle, preceded by a fuscous costal dot; cilia grey. Ilindwmgs 
and cilia grey. 

North Queensland: Cairns in December. Queensland: Redland Bay> near 
Brisbane, in November. Two specimens. 

Cen_ Euageta nov, 
ivayyTos, clear bright. 

Palpi long, porrect. Forewings with 4 and 5 stalked, 8 and 9 stalked. Hind- 
wings with cell long, 2 from beiorc angle, 3 and 4 stalked to near tenncn. 
5 absent. Distinguished from Euzophera by the porrect palpi. 

Euageta arestodes n. sp. 

('/tetrrwSys, pleasing. 

t? , V. 14-16 mm. Head and thorax pale fuscous. Palpi 4; grey, base, 
beneath white. Antennae and abdomen grey. Forewings with costa straight, 
apex rounded, tenncn oblique; pale fuscous; a white costal stripe from near base 
to near apex, broad in middle, narrow at each end; costal edge pale fuscous; 
a slender or suffused white subtennirud line; some minute terminal dark fuscous 
dots; cilia white. Hindwings grey-whitish: cilia white. 

Queensland: Tnjune in April (W. B. Barnard) ; two specimens. Type in 
Queensland Museum. 

Gen. Catkremna Meyr. 
TTdbk. Frit I.ep., 375. 

Palpi ascending-, recurved. Forewings with 4 and 5 separate, S and 9 stalked. 
Hindwings with o and 4 stalked, 5 absent. Type. C, Urcbdla Zinck., from 


Oteremna microdoxA Mcyr, 

Proc. "Linn. Sot\ N.S.W.. 1880,231. 
Darwin, Duaringa, Launceston. 

Catkkkmxa ati*rpnks Turn, 
Froc. Roy. Soc. Qlcl., 1912. 125. 


Trans. Roy. Sue. Qld., 1912, 125. 
Stradbroke Island. 

Cateremna QUADKiGurnaxA Wlk. 
35, 1,711- 

1348 mm. Head, palpi, and thorax fuscous. Antennae grey. Abdomen 
grey, bases of segments .sometimes fuscous. Forewings rather narrow r , posteriorly 
dilated, costa almost straight, apex obi use, tcrmen almost straight, slightly 
oblique; white with fuscous markings; a broad oblique bar from base of costa 
to dorsum, a bar from one-third costa to rnid-dorsum, where it joins a large 
suffusion broadening to termen and connected to apex; two blackish dots placed 
transversely in disc at two-third? ; a slender subtenuinal line edged posteriorly 
white followed by some grey suffusion, a terminal scries of blackish dots; cilia 
vhirish. Hindwings grey; cilia whitish. 

Darwin, Cape York toLismore, Milmerran, Mount Lofty, Western Australia. 

Cateremna cataxutha n. sp. 
KaTo£ov0a?, tawny posteriorly. 

3, 9, 14-16 mm. Head, palpi, antennae, and thorax grey. Abdomen 
fuscous; apices of segments and tuft pa!c ochreous. kegs fuscous with whitish 
rings', posterior tibiae whitish. Forewings with costa moderately arched, ape* 
round-pointed; termen straight, slightly oblique: grey-whitish sprinkled with 
fuscous; a slender median dark fuscous line forming the anterior margin of a 
broad transverse fuscous fascia and preceded by a fuscous dorsal spot; a slender 
wavy fuscous subterminal line edged potcriorly with whitish ; an interrupted 
fuscous subterminal line; cilia grey, bases whitish. Hindwings pale ochreous; 
cilia grey-whitish. 

North Australia: Darwin in December. Queensland ; Toowoomba in 
October. Ten specimens. 

Cateremna mediolinca u. sp. 
■niediollnms, with central line. 

9, 16-18 mm. Head, palpi, thorax, and abdomen white. Antennae grey. 
Fomvingr> narrow at base, costa gently arched, apex obtuse, lermen slightly 
romided f moderately oblique; rale brownish-yellow; a median Avhite line from 
base of costa to termen above middle, its margins sprinted with blackish above 
and beneaih; an obscure white subdorsal line sprinkled with blackish; a narrow 
grey-whitish terminal fascia sprinkled and margined with blackish; cilia grey- 
wdiitish. Hindwings and cilia grc\\ 

North Queensland: Cape York in April and June (W. B. Barnard \ ; two 
specimens. T}T- C in Queensland Museum. 

Cateremna leplo^tila n. sp. 
A£7rrorrr/Afjj, narrow- winged. 

9, 14 mm. Head, palpi, thorax, and abdomen fuscous. Auiennae grey. 
Forewings narroAv, dilated posteriorly, costa slightly arched, apex obt^c, termen 


oblique; whitish sprinkled with fuscous; markings fuscous a line from base of 
costa to two-thirds dorsum; a postmedian square uniting this with eo=ta; a very 
fine line from costa near apex to two-thirds dorsum; some terminal suffusion; 
a terminal series of dots; cilia fuscous. Hindwings and cilia pale grey. 
New South Wales: Broken Hill in March; one specimen, 

Catebemma lettcarma Meyr. 

Proc. Linn. Soc N.S.W., 1880, 230. 

Brisbane, Cunnamulla, Sydney, Birchip. 

Cateremna albicostalis Luc. 

Proc. Roy. Soc. Qld., 1891, 93. 

Cairns, Atherton, Townsville, Mackay, Bundaberg, Brisbane, Stradhroke 
Island, Lismore. 

Cateremna metallopa Low, 

Proc. Linn, Soc. N.S.W., 1898. 46. 


Cater emna melanomita n. sp. 
jitcXairojUTOs, Willi blackish lines. 

o . 18 mm. Head and thorax grey. Palpi grey; second joint with post- 
median whitish ring. Antennae grey. Abdomen fuscous; extreme base of 
dorsum and tuft whitish. Forcwings dilated posteriorly, costa slightly arched, 
apex rectangular, termen almost straight, slightly oblique; grey densely sprinkled 
with fuscous to middle, thence slightly; a sinuate blackish line from midcosta to 
mid-dorsum; a similar doubly sinuate subterminal line; a terminal series of 
blackish dots; cilia grey Hindwings and cilia grey. 

North Queensland: Mackay in October; one specimen. 

Cateremna pamphaes Turn, 

Proc. Roy. Soc. Qld., 1904, 47. 
Darwin, Townsville. 

Cateremka iiemiuapiies Turn. 
Proc. Boy. Soc. Qld., 1904, 47. 

Sea Lake, Hobart, Quairading, Perth, 

Cateremna AroDECTA Turn. 

Proc. Roy. Soc. Qld., 1903, 129, 
Brisbane, Scone, Sydney. 

Catekemna thermocukoa Low. 
Trans. Roy. Soc. S. Aust., 1896, 160. 
Darwin, Hrishane, Cardiff, W. Aust. 

Catekemna odontoses a Turn. 

Proc. Roy. Soc. Qld., 1912, 126. 

Cairns, Imbil. 

Gen. TylochAkes Mevr. 
Ent. Mo. Mag., 1883, 256. 

Palpi ascending, recurved. Forewings with 4 and 5 stalked, 8 and 9 stalked. 
Hindwings witb cell short (one -fifth to one-fourth), 2 from near angle, 3 and 
4 stalked. 5 from angle. 


Ty loch ares cosmiella Meyr. 
rroc. Linn. Soc. N.S.W., 18/8, 212. 

Diuiringa, Brisbane, Sydney, Moruya, Broken Hill, Melbourne, Birchip. 
Mttrtoa, Wirrabara, Mount Liebig, Perth, Rottnest Island. 

Tylochares epaxia n. sp. 

cVttems-, handsome. 

9 . 23 mm. Head brown ■ face whitish Palpi and antennae grey. Thorax 
gtfeyj patagia whitish. Abdomen oehreons with a series of central fuscous dots. 
Fore wings with costa almost straight, apex subreetangular, termen slightly 
rounded, scarcely oblique; grey, a well-marked straight white .sub-basal 
line, followed by a broad fuscous transverse fascia suffused posteriorly: an out* 
wardly curved slender whitish subterminal line, indented beneath costa and above 
dorsum, edged posterior!)* by a .series of confluent fuscous dots, an interrupted 
fuscous terminal line preceded by whitish suffusion ; cilia grey, apices whitish. 
Hindwings bright oehreous; cilia grey, apices white. 

North Queensland: Lake Barriue. Athcrton Tableland, in January (F. J. 
LHunigan) ; one specimen. 

Tylchares prays n. sp. 
■npa'VK, gentle. 

S. 24 mm. "Head, antennae, and thorax grey. Palpi grey; apices of .second 
and terminal joints white. Forewings dilated posteriorly, costa rather strongly 
arched, apex obtuse, termen slightly rounded, slightly oblique; grey sprinkled 
with whitish; a fuscous basal patch containing an oblique outwardly curved 
blackish transverse line; closely following this a slender outwardly curved grey 
line from one-fourth costa to. two-fifths dorsum; a grey median dot above middle 
and another beneath two-thirds costa; a suffused grey spot above tiiree-hTths 
dorsum; a broadly suffused grey submnrginal line not reaching tornus; a terminal 
series of grey or fuscous dots; cilia grey; apices white. Hind wings pale grey; 
cilia white with a grey median line. 

Queensland; Bunya Mountains in January; Stanthorpe in March. Two 

Tvlochakes scepttjcha Turn. 

Proc. Roy. Soc. (3ft, 1903, 130. 
Stanthorpe, Gisbornc. 

Tylochares gypsotypa n. sp. 

Artf'OTvTros, with white marking. 

£ , 2 . 18-21 mm. Head, antennae, and thorax grew Palpi long, much 
exceeding vertex, in male 2 and a half, in female 4; grey, towards base white. 
Abdomen ochreous, towards base grey. Forewings narrow, costa slightly arched, 
apex obtuse, termen obliquely rounded; grey; a broad white stripe, sprinkled with 
grey, narrow at each end, from base to apex; sometimes i\ series of minute fuscous 
terminal dots; cilia grey. Ifiitdwings pale grey with darker terminal line; cilia 

Xorth Queensland: Cape York in October and November ( \V. B. Barnard) ; 
two specimens. Type in Queensland Museum. 

Tylochares anaxia n. sp. 
^a£ios, of little worth. 

$\ 5 ■ 16-17 mm. Head, palpi, antennae, thorax, and abdomen grey. 
Forewings dilated posteriorly, costa slightly arched, apex obtuse, termen slightly 


oblique; grey; a fine whitish transverse line at one-third, angled outwards in 
middle, preceded and followed by more or less fuscous .suffusion; short fuscous 
streaks on veins in terminal area; a terminal series of fuscous dots; cilia grey- 
whitish. Hindwings and cilia grey-whitish. 

North Queensland- Cape York in November, Dunk Island in May; lhree 

Tylochares paucinotata n. sp. 
paxtcinotatus, scantily marked. 

9 . 16 mm. Head and thorax fuscous-brown; face pale brownish- 
Antennae fuscous. (Abdomen mining. ) Forewings narrow, dilated posteriorly, 
apex rounded, termen obliquely rounded; greyish-brown with some fuscous dots; 
three placed transversely al one-third; a median dot; an outwardly curved sub- 
terminal line of dots; cilia grey, Hindwings grey-whitish ; cilia whitish with a 
faint sub-basal grey line, 

Cape York in October f\V. B. Barnard); one specimen, 

Tylochares ktckmoxoju* Turn. 
Proc. Roy. Soc, QUI., 1912, 125. 

Tylochares psoleuca Low. 
Trans. Roy. Soc. S. Ausfc, 1903, 58. 
Brisbane, Dendiquin. 


Proc. Roy. Soc. Old.., 1912, 226. 
Caloundra, Brisbane, 

Tylochares chicnopleura n_ sp. 
^toro7rAtv/>a5. with snow-while cosia, 

o,9. 16-20 mm. Tlead.palpi, antennae, and Ihorax fuscous. Abdomen 
pale nrrey. Forewings narrow, co^ta straight, apex rounded; fuscous; a snow- 
white costal stripe from near base to near apex; codal edge fuscous; cilia white, 
on toi-nus grey, but sometimes wholly grey. Hindwings and cilia pale grey. 

Queensland: Warwick in December, from larvae feeding in Acacia galls; 
two specimens. 

Tylochares endophaga n. sp. 
M'So^or/os-, feeding internally. 

$. 21 mm. Head, palpi, and antennae grey. Palpi not exceeding vertex; 
grey, towards base white. Abdomen with basal half ochreous, terminal half 
grey. Forewings. with eosta slightly arched, apex rectangular, termen straight, 
not oblique; grey; a white stripe, sprinkled with g-'-.-y. from base to near apex; cilia 
grey. Ilindwhigs grey whitish; cilia whitish. 

Queensland: Bribie Island, near Caloundra, in October; one specimen from 
larva feeding in an Acacia gall. 

Tylochares pastopleurs n. sp. 
TraiTTCOTrXcvfjQ'i, with sprinkled costa. 

c, 5. 15-16 mm. Head, antennae, and thorax grey. Palpi fuscous. 
Abdomen whitish, in female slightly orhreous-tiuged. Forcwmgs narroAY, costa 
slightly ached, apex rounded; grey; a costal stripe from base to apex, over one- 
third breadth of wing, sprinkled with fuscous; a terminal series of minute fus- 
cous dots; cilia whitish or grey- whitish. Hindwings wild cell one-third; urev- 
whitish ; cilia whitish. 

North Queensland: Cape York in October and November (W. B. Barnard) ; 
two specimens. Type in Queensland Museum. 

Tylochaues goxiosticha Turn. 
Trans. Roy. Soc. S. Aust. 1915, 803. 
Musgrave Range. 

Gen. Pkjipelia Hb. 
Verz., 369. 

Palpi ascending, recurved. Forewmgs with 2 from well above angle. 5 
separale, 8 and 9 stalked. Hindwings with cell short (one-fifth), 2 from angle, 
3 and 4 stalked., 5 absent. 

Peiipeeta opimella Meyr. 

Proe. Roy. See. N.S.W., W78, 201. 

Brisbane, Mount Tamborine, SLauthorpe, Miles, Milnierran. 

Pempelia caniljxea Meyr. 
Proc. Linn. Soc, N.S.W., 1878. 209, 

Brisbane, Toowoomba, Carnarvon Raivrc, Mirrrmundi, Sydney, Goulbtirn* 

Pempelia iio-ncuLAENA Meyr, 

Trans. EnL Soc, 1887, 260, 


Tnm B . Roy. Soc. S. Aust... 1893, 166. 

Gen. Tiussoxca Meyr. 
Proc. Linn. Soc. N.S.W., 1882, 158. 

Palpi ascending, recurved. Forewings with 5 separate, approximated at 
origin, 8 and 9 stalked. Llindwings with cell shorl (one-fifth to one-third), 2 
from angle, 3 and 4 stalked, 5 absent. 

Trissonca clytopa n sp. 
k\vt(o~(is, noble. 

t, 9. 14-17 mm. Head fnscous; lower edge of face whitish. Paipl and 
thorax fuscous. Antennae grey. Abdomen ochreous. Legs fuscous with whitish 
rings; posterior pair whit"sh. Forewings rather narrow, posteriorly dilated, costa 
straight to middle, thence arched, ternien stai^ht, scarcely cblkrcc; a fuscous 
basal patch limited by a sleuier IdackLh line posteriorly from two-fifths cosla to 
mid-dorsuiu. angled outwards beneath costa and ^Jtiftt above middle; disc beyond 
rh : s rrroy or partly grey-wln'tish, with some fuscous admixture; an outwardly 
curved blackish sttbtcrminai line; a slender blackish subrnargiual line preceded 
Ly more or less white suffusion; cIHa r>Tey, apices whitish. 

-North Queensland: Kuranda (V. P. Dodd) \ two specimens. 

Tkissonca iaktiiemjs Meyr, 
Trans Lot. Soc, 1897, 250; ep&t&fnvs Turn., Proc. Roy. Soc. Old., 1904, AS. 

Darwin, Cairns. Atherton, Townsville, Bp^Mtj F.uiigdla, Yeppoon, Brisbane, 
Toowoomba, Charleviile, 

T^tssunca moia ronton a Low. 
Trans. Roy. Soc. S. Aust., 1903, 5/. 

Cooktowu, Towr.svnle, Xambour, Brisbane, Tweed Heads. 

Trissonxa i-iesactella Meyr. 
Proc. Linn. Soc. N.S.W., 1879,225. 


Gen. IIypocjryphta 'Rag. 

null. Soc. For. Fr.,, 1890, 119. 

Palpi long, povreet. Maxillary palpi iti male penieilJate. Forewings with 
4 and 5 stalked, 8 and 9 stalked. Hind wings with cell short, 2 from angle, 3 and 
4 stalked, 5 absent. 

Hypogryphia amicto&cs n. sp. 
au^Tw?-)?^ unmarked. 

EJj 9. 22-23 mm, Head and thorax greyish-Vo--^. Palpi 4; grey, near 
base w! itisb. Antennae and abdomen grey. Forcwings narrow, posteriorly 
dilated, costa sa-aight to middle, thence genttj* arched, apex rounded, termer 
obliquely rounded; greyi sh -brown ; cilia eoneolorous. SfilidwtDgs pale ochreous- 
grey; ci-ia whitish. 

Queensland: Duaringa ijy December; Toowoomba in October (\V. B, Httr- 
nard) ; two specimens. Type in Queensland Museum, 

Gen. Ancylcsis Ztk 
L?is, 18e9, 178. 

Palpi porrect. Maxillary palpi hluorm. Forewings with 5 separate, 8 and 
9 stalked. Hfndwings with cell sbort, 2 from angle, 3 and 4 stalked. 5 absent. 
Type, A, cnihamomca Dup. t from Europe. 


29, 829. 

c , 9 . 14-16 rnni. Head, palpi, antennae and thorax fuscous. Abdomen 
pale grey. Forcwings with costa sligfetly arched, apex round-pointed, iermen 
straight, slightly oblique; pnle oehreot;;>-grey ; a fuscous do - , below middle at one- 
third ; a narrow dark fuscous terminal fascia; cilia grey ; apices whitish. 

North Queensland: Townsville.. Oueenslaiid : Yeppocn. Also from Ccylom 

Ancvlosis kctifascia Pimps. 
Rom. Mem, 8, 193, 

pi $. 22-25 mm. Dead and thorax whi.tish-oc.hrcot $ ; patagia and tegttlae 
pinkish. Palpi 6] lower edge whi:e. Antennae pale grey. Abdomen grey* 
whitish. Forewings who costa straigh: to mkld'e, thence arched, apex pointed, 
terrncu very obliquely rounded; whiush more or S053 pinkish-tinged with slight 
gn:y Sprinkling; a median line from base In two- birds or more, edged nlxue with 
white, beneath with blackish scaler; a terminal series of blackish doirj ; cilia 
whitish. Mindwings gri;\-whu : sh; cilia whitish. 

.Xor:b Queensland : Towusviitc. Queensland : Peak Downs. Gayndah, Natn- 
bour, Er'shaue, Dalby, lnjune. New South Wales: Scone. 

AncyloSi'3 thf.ostichn n. iyj. 
Ouoirrt-xo:, M_dphur-l;ucd. 

8 , g\ 22-24- mm. J lead and thorax ochEOous-g'rcy. Palpi 6; grey, hwer 
edge except terminal, ioini whitish. Antennae grey. Abdomen fuscous. Pore- 
wings with costa gendy are-bed, apev obtuse, icemen ohiovWy rormded; grey, in 
male sa:ffused with whitish; in female a fine pale yellow hue from base to apex; 
near apex, a broader stihmedian Ffntf from base to tenr.en; a similar subdorsal 
line much narrower except near base: a subcostal fuscous dot at onothird and 
another at end of eel 1 ; a termira! seres of blackish dot:;; cilia grey; li hid wings 
grey; cilia grey, towarJs dorsum whilish. 

Queensland: limine in October (\V. B. Parnard ) ; two specimens. Tvue in 
Queensland Museum. 


By Keith R. Miles, Department of Mines, Adelaide 


A review of the Australian Pre-Cambrian Succession reveals occurrences of granites and rocks of 
granitic appearance encountered over very considerable areas and apparently involving vast periods 
of geological time. With the changing modern concepts of the origin of granite and of granitic- 
looking rocks, all evidence concerning the relationships of these rocks in both time and space, both 
with one another and with other adjacent geological formations, assumes a new significance, from 
the point of view of both historical geology and the specialized problems of petrogenesis. It is time 
that such evidence should be brought forth once more and critically re-examined in this new light. 
Periodical stocktakings are always valuable, and, to the scientific worker, can prove both salutary 
and encouraging. 



J j y- Kcmi R. Mjles. Department of Mines, Adelaide 

[Read 14 November 1946] 

Plate I 


Txmmurnr>:>' .. .. .. ,. .. .. . . 54 

Qkigtn OS GkAmtf. — M'.Hir.RV CVjvcr.pTs .. . - .- . . ..54 

GAwmsATrax .. r i .* .» .. .. „, . * 53 

(JRAN.tTISA-*rOM AMI THK PeK-CaVHRIA-C Si TCrJS.STiJX tf .. .. .,57 

W'kstfrn Arsr.tALiA ._ .. .. . . . . .. 5S 

Sol rn Alistuaua. , t .. . . .. , . .... 59 

Ixu;\'!jmic Qwsrnru.vnoxs . ► . . . . . . , . 63 

Co\ t (i.v^ok .. ,„ .. .. .. .. .... 6-1 

Ac K:-;o\vi.r.;>t;\iF\'T:? ,. ,, .. .. .. . .. 65 

A review of the Australian PrMJambria.n Succession reveals occurrences of 
granites and rock? of granitic appearance encountered over very considerable 
area** and apparently involving vast period s of geological time. With the chaii£- 
itig, modern concepls of the origin of granite and ot granitic-looking rock?, ail 
evidence concerning the relationships of these rooks in both time and space, both 
with one another and with other adjacent geological formations, assumes a new 
significance, from the point of view of both historical geology aud the specialized 
problems of pedogenesis. It is time that such evidence, should be brought forth 
once more and critically re-examined in this new light. Periodical stocktaking? 
are always valuable, ^nd, to the scientilic worker, can prove both salutary and 

Processor II. II. Read in two memorable presidential addresses to the British 
Geologists Association, emitted "Meditations on Granite, Parts I aud IT" (Read 
1943. 1944). has priced all Engiish-speakinjLr geologists forever deeply in his debt 
for a clear presentation of the facts of ihe problem of the origin of granite and 
of the modern trend of thought, which are thrown into proper perspective by an 
exceed tngV ilhiminaling historical background. This historical review, like atl 
good science, has a truly international flavour and follows the arguments and 
ec:r l n'o\e/bies of the P»rilinh v French and Fennoskanclian masters from the kit:? 
18th century up to the present day. It has probably come as a -hock to many 
Australian geologists to learn thai ideas very similar to the present conception 
of "granitisatioi-T were first put forward by the Frenchman (Ami Bond) in 1824, 
and that Hie thesis lias been steadfastly developed and elaborated by most of the 
STeat French g-ecMic^sts from thai inue onward. Such ideas were diametricallv 
opposed to the principles of igneous geology laid down by leading penologists of 
the German School, under whose Tar-reaching influence British and American 
thought, uu ' 1 ely. remained clouded until well into the 20th century. 

The old concept of "ieneous" rock based on the fundamental three-fold 
classification of rocks into Igneous, Aqueous or Sedimentary, and Metamorphie, 
the first rung of the ladder to which most of our first student steps were guided, 

Trans. Roy. S»o S. Au&L 71 (1>> £5 Jul.* 1947 


is proving untenable in certain important respects and can no longer fully sn.-tain 
us on our upward climb. This fact has been amply demonstrated by Read ia 
his review of French and Fcnuoskandian literature and supported by ihe latest 
researches of many workers in both Britain and America. It has become increas- 
ingly apparent thai there is a fundamental genetic difference between ibe two 
main groups of so-called "igneous 7 ' rocks, Lc, basalt and granites, and that con- 
trary to past belief* an understanding of the origin* mechanics of emplacement, 
trend of differentiation, by-products and so on o£ the first .croup in general con- 
tributes very little towards the understanding of the same features of tbe latter 
group. YV. Q. Kennedy (in Kennedy and Anderson. 1938), discussing this 
problem, has advance J tbe idea of two apparently independent expressions of 
"maprnatic" activity called Volcanic and Plutonic Associations-— the former con- 
sidered to he derived from a universal basaltic magma, winch has originated from 
the renieUins* of a basaltic eartli shell, the intermediate layer. The latter are 
considered to be derived from a primary universal granoiioritie parent magma 
which bus developed by remeluug oi the -so-called '^Granitic' 4 \$y& within 
orogctiic zoned, where tectonic thickening at ihe earth's crust has brought the base 
of the granite within the range of meUirur- Important differences in the mode of 
irruption in the two cases have been cited. The granite and granodioritc bathohihs 
appear to penetrale slowly upwards, accompanied by a wave of granitisation and 
migmatisarion of the country rocks, until arrested by some unknown form of pre- 
sure balance akin to hydrostatic control before they reach the surface. The asceiit 
of basaltic magma, however, is believed to be directly towards the surface by wcy of 
a system of relatively narrow dyke-like fissures with no large intercrustal reservoirs 
being formed. Tbe magma is then ci'her extruded as lava Hows or forms injected 
bodies of various sizes, such as sills or laccoliths, which may themselves represent 
volcanic reservoirs. IS'o large scale migmutisation or metasomatic replacement of 
country rocks is ever performed by sueh basic, magmas, whose differentiation and 
subsequent ev'uiion arc controlled largely by fractional cry* Utilisation processes. 

"Without necessarily conceding the validity of all of Kennedy's conclusions 
vj their entirety, it js apparent that there are certain majt r irreconcilable differ- 
ences in the mode of occurrence of these two groups of rocks pointing' to two 
quite distinct forms of so-caked "igneous v activity. This is nowhere more clearly 
demonsi tabic than front studies oi Ihe Archaean granitic rocks ihroiighoiu the 
world, from which it is also b.'roming more, and more clear 1 bnr the- div-siou 
between "Igneous" and "Meiauiorphie" in the old three fold classification, u\. 
as far :js Kennedy's "plutonic association'' of granitic rocks and tbe high-grade 
mctamorphn- rocks are conn-mod. is more apparent than real, and lhai in ninny 
cases die distinction between orlho- mm\ para-gneiss completely breaks down, \t 
is from tbe observation of insiauee- ithisi ruling this l;i>: farr and by deduorions 
therefrom that we have reached this modern viewpoint on the origin of granite. 


Tho pub oi* this viewpoint can probably hest be summed up in the word 
"grnmhsaiion/' a word which according to Grout (1941 "1 has been loosely used 
bv writers for a number of years and which has recently heen defined by Read 
£ 1-9441 1 n.ttcr an analysis of definitions from many soir/ccs, s& "the process by 
winch solid rocks are converted to rocks oi granitic cha -aeter without passing 
through a magmalic stage." Some h'reneh writers apparently Hft£ the term 
fplbpalfi' l .adou as synonymous with granilisa'don. 

Gr;uiitvs;ition of country rock is dependent upon the "nlroductiou and mela* 
somaiis-hig action of some form of v?r\ active emanations -gasemir., fluid or both — 
whose origin is yet obscure Vtase rmanathms, whetfjer vapours or solutions. 


ajn* apparently extremely active and tenuous and must be at least heated above 
;he critical point of water ,( '365** 1 L\ at approximately 200 atmospheres), liu* 
mechanics of introduction of these j*fran$WrMlg emanations have been discussed in 
considerable detail, by Prcnch ami Pcnuoskandimi authorities. Apparently two 
ehie! processes are recognised — lit par lit injection or preferential replacement 
ui:d n form of hodily (jfapnijoal replacement through "soaking up" or "imbibition"' 
(to nse the Prench term} without distortion or displar.Mnen: o(* the counlrv rock, 
and migration of materia] Lry what has been termed "oil ?pr!" mechanism. Jn tins 
respect C ft. Wegmauii 1 ,1931, 1935. 193 S) has emphasise- 1 the importnucc of 
inrer^ranuiar films in tbo (Molecular replacement and uvirrnt'oii of material -and 
similar reactions. 

One of the BioSt titt&difcjj features of granirisation is ffhy ajid how the iinal 
result of the permeation and replacement process is apparently always the forma- 
tion of granite or granitic, rock, irrespective of tbe composition of the country rock 
attacked. Tin'.- has been explained in the concept of migration and advancing 
'•froins/' developed by Wegmann (op, at-), ttackhuid (1938), Reynolds (1^43, 
1944) and others. Material from the gratniising emanations is regarded by tbeoi 
$$ selectively replacing the components of the pre-existing rock, adding hc:re v 
taking away there, so a? to leave remaining" materia! granitic in composition. The 
displaced material is driven forwarr! with outgoing emanations and fixed in .an 
outer zone or "front." The geochcnncal relationships involved in granitisation 
have been expressed in mathematical terms by Holmes (1945) as: "Granite == 
pre-existing rock plnx added material (A) introduced by and ahstraetcd fiuin 
incoming emanations (A -j x) minus displaced material (til driven forward with 
outgoing emanations (B -f-x). 1 ' The remarkable feature about these cmanatiorii- 
(A -J- x) is that (x) apparently leaves no trace, from which we may deduce 1ha: 
they cannot represent ordinary ina^ma. 

Qualitative geuchemicat suirb'es of toe successive stages of e;rankisation in 
the field recetitly carried out by !)r ( Doris L. Reynolds on the Ne\\r> Igneous 
Complex 1 194,5, 1944} have demonstrated that the minimum imroductions (A) 
were Xa. Ca and Si, and displaced materials (H) eveniual!v carried forward wtrt' 
Al, h\\ Itoj, K, K, Ti, P' Mn. These (predominantly basic J displaced materials 
were found to have been fixed in an aureole of "igneous Inching basic imd uhra- 
basic tucks, and it is crmchidcd from tfrcsA studies that befoie any given mass <>t 
eoitnrn rock was granitised ir passed tftiVUlfth a preliminary stage oi basification 
In a very recent paper (1 ( )46) (abstract only avallab'e ai time of writing) Or. 
UeuolS has: enlarged on this subject and demonstrated that the geocheinical 
changes leading to granilisation in rocks of all types invariably includes an mit ; a! 
enrichment in Femie constituent* aivl alkalis ("hasiricatiou, 1 ' "dedication") Vol* 
lowed by a stage ot fe^pruhisatiou or .grunitis-rr.iun proper. Details of the stages 
in nietasornatje. a'tcralion of pe'itic, seini-pehre. psummitie and fra&tc igneous 
roH-:s .are given, providing valuable criteria for die recognition of the provenance 
<»f sWiocths or "enclaves*' in granm*. 

Ill considering the origm oi granite perhaps the most satisfying feature of 
the "replacement metamorph:?m" hypothesis is in connection Avith the space 
prohhan. particularly as it concerns the enormous regionally imeissose granitic 
Uias-cs of the Archaean sfrehU and ancient continental blocks ami the major 
h;onoi!tb* usually torming cores* tu fo'ded ranges of periods later than Arehfien- 
y.oic. In horh these ca?^ if Ike granite is considered to be .strictly iu'msivc, that 
k. add* -I to Ll:e pre-existing rocks, then die .space factor renders this view 
mitcnaLle. There* is no ev;rkuec of dlspkicemcnt of country rocks on even I lie 
•smallc,-; irn.iiuu oi the $czx remiircd, nor has there vet been imagine! auy 
meLbani'*m ai rtdituttjj, subsidence, block tanltiu^, -,topin;x/etc., t\y a ; couLu accuulil 


for the entrance of such volume of magma Involved. Ail the evidence, on the 
other hand, points to 3rQ>JscfUlCt>l vyult little or no bulk changes in volume. How- 
ever, as far as minor granitic intrusions such as dykes, sills and v:*lns of any age 
arc concerned, the actual injection of material as liquid, i.e., magma, inio places of 
weakm-ss would appear to be an equally logical explanation. 

Finally, attention may he drawn to the division of granites into the three 
categories enumerated above-, vis. : ( a ) Archaean graniti: masses, ( h ) core 
batholiths of later a^c, and (c) minor intrusions of a!l aires, and to the sugges- 
tion, tol* which there appears to he some claim, that there was greater igneous and 
mctaniorphir activity in Archaean limes than at any later date, with the inference 
of a possible fundamental difference in origin between the Archaean metamcrphic 
rocks (particularly granitic) and those of later periods. 

We lviaj now come to a consideration of the relationship of the graniiisatioa 
replacement concept of the origin of granite to the question of the Prc-Cambrian 
Succession in Australia. In the light of the modern viewpoint, i: is necessary to 
re-examine evidence as to relative ngj of granite emplacements and the petro- 
gencsis of adjacent rocks- II is now clear that apparently petro^raphically dis- 
similar types of granitic rock, c.y,, an apparently missive granite, product of flic 
granitisation process, and an adjacent granitic gneiss or schist, product of partial 
irraniiisafon, and also possibly a more basic type of igneous looking rock — say 
dioritir or mnn;>rmitic. representative of Wcgmamvs Mg-Ke ''front'— may all be 
elosc'y linked both in age and orkpn. Furthermore, it is evident that pressure 
movement* resulting from adju^ment of minor volume changes in the country 
rocks during I he granitisation process might result in bod ly displacements of the 
already formed plastic or fluid granite mash producing in* aisions of massive non~ 
gnc&bit granite into its own contemporaneous granitisalion products. 

In sueh a case it appears conceivable to the present writer that, especially 
where the granitiscd sediments show rapidly varying character and composition, 
the chemical composition and pctrog rap Ideal eharaeters of the intrusive- granite 
an 1 of the mure granitic portions of adjacent migmatites might vary to such an 
extent as to give the impression thai the two "granites" were neither con- 
sanguineous nor contemporaneous. It is possible, therefnie, that limited field and 
laboratory observations may have ui some, cai-es led to entirely erroneous con- 
clusions as to the relative age of granitic magmas. Further, if the granitisation 
ther.ry [3 accepted, then the correlation of granite on chemical and mineral com- 
pnshion alone obviously becomes increasingly difficult. 

Consequently it is opportune to review such evidence as is available recording 
lite periods qf emplacement of all bodies of 1 're-Cambrian plutonic rocks, both 
the grannie and the more baste types, in the. hope that a elenrrr picture may 
emerge of the nature and scope of Prc.-Cambrian igneous activity throughout the 
Commonwealth, It it considered that by making studies of Lhis kind, interesting 
evidence should be obtained to aid the elucidation of some, of the mysteries of 
the Archaean basement complex, ihe primeval foundation upon which ail other 
geo'o^irul formations have subsequently been built. Ii i: to be anticipated that 
Mich w-'M'k will also lead to some inte-esting conclusions regarding relationships 
in ue deposits to igneous activity and the granitic rocKs t and will thus have, con- 
siderable economic significance. 

'J 'here is no doubt that there arc in Australia mimerou-i areas which would 
provide excellent scope for studies of the phenomena of granulation and granite 
em]rlacement in rocks of many different uge.s, Some 01 the^c occurrences were 
flescrlbed and discussed at the meeting of A.N.Z.A.A.S. he'd in l^.te), but (lie 


pttbii&hfcd abstracts of papers submitted (Report o[ 24th Meeting- A.N\Z.A + A,S.„ 
Canberra. 1939, p. 95-98) shows that the conception of grauitisation held at that 
rime ddfered in several important a.speets from the more modern ideas already 
cxpre c -i\']. The terms ''^jtftfes&io** 1 ' and ''assimilaiioii" were then apparently 
used. In describe rather similar processes lakiug place at the margins of and 
consequent upon granitic magma intrusions. No clear conception of "the meaning 
oT "granuisatioiv' in its modern sense, or of its probable role in granitic magma 
format : on was then indicated. 

The " Fundamental Complex'* rocks — the granites, gneisses and lui^Tnatito' 
associaied" with high-grade schist? of the Archaean Shields — provide exposures 
cii the largest scale that are most suitable for these investigation-. The greatest 
area:! distribution of these rocks occurs in Western Australia, South Australia, 
Northern Territory and probably Oiieenslaiuk AUhough many such areas are 
situated in geographically remote regions and in consequence have not received 
detailed study, there are stdl numerous occurrences in reasonably accessible 
localities which are relatively unknown. The present writer's own persona! experi- 
ences are concerned with areas in Western Australia, and to a lesser extent in 
Sottih Amn-aha, which are the subjects of tfe paper. The notes presented ?H 
the following sections refer to observations made in studi areas, and are confnr -I 
in particular to ihe IVe-Cambrian granites. 


A glance at David's Geological Map of rhe Commonwealth of Australia 
rcvenls 1 1 to southern half of Western Australia depicted as composed very lar^ely 
of ''granite cd May-juno Creek Series, of Older Proiero/.oie Age." which enclose 
scattered "islands'' q[ metamnrphic sedimentary and igneous rock* representing 
the Yiltfarn and Ktffronfjfe Series, disposed in a general meridional tzfnd: The 
'granite" areas cm this gxttfti Archaean Shield ru reality comprise strict! \ limited 
outcrops of a wide and diverse variety pi acid mctamorphic and igneous took 
types — massive granites, banded or gueissie <r;-;m:tes showing every gradation r rom 
slight linear arrangement of minerals through more striked types enclosing relics 
of absorbed schists to obvious hybrids, composite gneisses or uiigniiitites, and to 
?ch:st> in which only minor quantities of interlaminaic I l %n£#tt5*' material em 
be distinguished, Lanjfe tracts of counlry intervening between these '"gramfc** 
oulcrops. particularly in the central and eastern ^oldlields areas, are covered with 
superficial deposits of sand, forming low. bread -y' undulating scrub-covered 
p]:-;iiis hi which outcrops are rare and comprise low. bare mounds or flat floors oi 
tna^swe granite. The rocks underlying ilwst? sandy plains are generally accepted 
as lidtig granitic or gmdssfc in e'-aracier. and are ofien distinguished in local 
mapping by tlui .-\mbol Gr/Gm (H, A. Ellis 19A t: i). 

I'l Sfi&tl3 clear that tins gfiftfr granitic massif, which can be reasonably 
;;, sinned io extend discontinuously over an area of the order of 200.000 square 
miles, cannot be regarded as having the generally accepted form of a batholith, 
rnr yet that of a series of batholiths, as this would imply a body or bodies extend- 
ing !o d<'p:hs of many hundreds of miles into the earth's crust. Rather, it would 
appear lliut it may have the form of a ,e;reat crusral sheet or sheets of considerable 
horizontal extent, but relatively shallow depth. 

The regional geological structure in s-nme of the "islands'" forming pwfffoto 
tff l!ie Western Australian goldnelds with winch the writer is familiar, z'ta.j Sic 
Y i ■ •_ ; ; ? 1 1 1 ( Ellis 1 9.30), jiortious of Cooigardic. \urrh Cuo^.olie ant .Mot;;.! 
Marv.nrct Gnle. field appears to be relatively simple, comprising in main ca.-i - 
comparatively gentle Infrtftrg w ah LjU& ilVic&n a that the pre-eni remnanis r+f t'a 
Kal^oorlie and YHgarn Series are the roots of original fold mountain chain; 


or the products of such oilier major otogenic movements as might be expected 
to accompany the bodily intrusion of vast masses of magma. 

On the contrary, generally .speaking the banding of the granite-gneisses over 
considerable areas coincides perfectly wi;li the regional and local struct urr.s found 
in the schists of adjacent ■i$I;tnifci' i of Yilgarn rr Kalgoorlie Series- whilst on a 
small scale banding- is frequently to he seen reproducing the structure unes ui 
original schists, of which relicts are sti'd preserved uureplaced. Jt is quite Apparent 
that the^e granitic rocks have not developed by the forceful intrusion and upward 
sloping of a granitic magma, but rather by some process of quiet soaking and 
replacement of the original country rock schists, t*i.::.. "gnmitisation." in which 
original trend lines were fir&t undisturbed, but as the ^'grar iiisatiun from*' moved 
forward llu: heated mass became, more fluid, pasty and probably culminated locally 
jll mngmn which could remain in place or move bodily n< an intrusive: mass, finally 
crystallising as massive granile such as that whose otu crops are round at ibfi 
isolated *'rocks*'* wuhin the tandy plains mentioned above. 

Some excellent exposures for the detailed mvestigaimu of the products of 
granitisation are ro be found in these yoldilclds regions, more particularly amongst 
the high-grade schists of the \ Again Goid field in such loralilies as Yeilowdinc- 
Mount Palmer district, 24 miles east rf Southern Cross, and the Hope's iiill- 
Coriotlnan district, running north from Southern Cross. An interesting feature 
of lbcse and other areas of the Western Australian goklhelds is the fact that a 
grent proportion of the original rocks arc basic in character— the Kalgoorh'e- and 
Yilgarn ''greenstones'' contain great Lh'xknesses of bade, probably mainly bas:dtic_, 
lava flnws metamorphosed in varying degree, and some bodies of basic and ulini- 
bas'c intrusive rocks, do'erites and gabhros. In many areas these rocks, rich in 
J r e and Mg, are now represented by and amohihohte schists. It is 
obvious that during regional granui-aiiou tit the original basic rucks enonnous 
quantities of Fe and Mg nrtut have migrated out of the rocks undergoing 
alteration, ;\-\m[ it i.s possible tbat many of (lie existing amphibolies ;r>" I amphi- 
bolitc schists may represent in fact the fixation of cafetruc material m a basic 
M fVoriJ preceding granitisation as has been outlined above*. These amphdiolites 
conlcl ihen perhaps he more accuraicly classed as aniphihoiiuc diabrochiivs (after 
Dunii, J. A., 1942). Similarly it wotrd seem possible that KlhV Greenstone Series, 
snhi'livisi ni of the. Yilgarn System, mJfry veproeir: hnl a broad "■front" nf bastrka- 
lion of the rneks which may have hal . n .n original composition similar to his 
WhiUMone Series. 

Derailed investigations of areas in the Darling Ranges near Perth, in recent 
years by P. % l 3 rider ,( 1941, 1945), and others, have resulted in an accuvnul-Uiuii 
of evidence for at lere-i two phases of Archaean granite emplacement in this 
region; (a) development of hybrid granite gneisses ;uid migmatUes from 
grauilis.Uion of pre-extsiing formal ions I possibly pari of the predominantly mcta- 
sed'inenlafy Cluttering-] impcrding Series, relicts of which arc exposed to the 
north-east til Perth ) ; (b) emplacement of a younger nuemdine granite which 
has engnlfed considerable portions of (a). The younger granite is considered by 
Prider lo belong ''lo a distinctly later period than the granite which gave r'--e to 
the lnhrid gran'to. gneisses." However, remnants or xennurhs of the original 
"bascmcnC rucks, comprising Pc-rich horiiblende-hiotite eoijote hornfels enclosed 
hi the granite gneisses which Prider considers Lo he derived from prc-exi-thtg 
basic n/neoos rock, may well represent 'Vhabro chutes," rc'iels of an oriruna! M;> Pe 
"froiu. 1 ' and under existing concepts of the processes of grau'tisation reviewed 
above it would, not appear inconceivable that the granite gneisses and the younger 
granite arc tnore cJoselv reined m Mge and pet'v-.'.renr^s than fcfes hi. herru I.kcu 


Aaother area of particular intercut which Prider lias recently described 
(]':; \r 1945b) fa ni Dan^m, 90 nriU-s cast of Perth, where lenticular xenolirli.s 
of rkaruockuic rock types — rang jug- from ultrabasie (oil vine-hornblende and 
eordieri'te hypersthenic*) to haste (p!agiorlascdiornh\m5c-pyro\cne eranuHtes) — 
an* found in country rocks of hybrid gneiss or "acid eharnockitos ,t 
(ccndi 'riie-hypcrslhenc-q'uartz-fcl^ gneiss, etc. ). PiTVr eonc'rides that the 
a:: ; d t.-r~rnoekires have de \ tdo]iC".i by ^ranh^aiion of prr-exdstm^ hypersthcuc- 
ben^rjn; basic chai*uoc.kties whu-H are ihem^elve^ the rc-crystallised products of 
hi±yc rvi'.rusions contaminated by asdmdalion of aluminous sediments. Here 
a<.;a : i.\. by aup'ieation of the theory of a Inincing* fronts, it would appear that the 
devf-jomout of the ultnibaste and basic charnoekites may have been a necessary 
pre'mvinary step towards the graniti.-adon of the original basic sedanenrs to form 
acid chanioc-kite tmehscs and may falls have &aeU an integral part pi the one 
pmcc.-.-. Thc.^c occurrences rtre of particular interest to us in South Australia 
lif.cau. >• of the development of Sifsnifo* rock typft; amongst thz- granite gneisses tfl 
liyro Peuinsu'a or'gimilly (inscribed by TtTey (1921). 

£ Uhcr areas iij Western Australia where excellent exposures of Archaean 
rrrarite and trneissvs suitable fur iniensive investigations are to be found arc in 
llvj h~t of "peripheral gneisses" extending around the coastline from Cape 
Naturalise in the south-wesi corner of die Stare to Albany, and thence 
eastwards for many nfJes along the south coast, and it is hoped that these studies 
will he* made in the light of recent developments concerning the origin of granites. 


"Lore it] South Australia contributions towards the r>etroe;enesis of the Pre- 
Lauii'.' can granitic rocks have been disappointingly few. Tins can only partly be 
exp_umi j d Ivy the relative inaccessibility of many of the more interesting outcrops 
of thesis rocks. Rather it would appear that sJttdic^ of the stratigraphy of the 
rroie-ozoic formations have completely over-shadowed consideration of the older 
"tundauietilar' rocks. 

RYenianT (1937) investigation of the 'Thorite" inclusions in the granite of 
Grairhe island pointed towards lite possibility of replacement of intruded country 
rock to form granitic rock, whilst AldertnanV. (1938) study of the chemical and changes involved in the production of the augen gneisses of the 
llumhuv; Scrub area •stands out as a notable contribution to the pedogenesis of 
these rocks. The augeu iujcc:ioTi gneisses and pegmatised schists of 
Uvfe region art.* considered hv Alderman let he due to intimate injection or permea- 
tion ot original sedimentary schists ( ?phyiihes now sericite schists) along bedding 
rend foliation planes by alkali silicate solutions. This is apparently conceived £ts 
an additive metasomatism iv, which it is calculated thai volume increases up to 
approximately double the original volume woiud be incurred during the reaction jt. 
lnu-rmd -tresses set up in the ,gne : s.-v.- as a result are concluded by Alderman tO 
have L'een responsible for subsequent dynamic meiamorphism of the rocks. 
V'ni'oi innately the value of this work is weakened, in the present writer's 
op-'nion. by Jack of supporting field evidence. When the considerable body of 
tfchfeU and injection gneisses in the whole area is considered, volume increases 
of the order indicated might he expected to produce more noticeable effects than 
mrre local granulation or mylouidzation /fines within the schists. 

.Rock exposures in. portions at least of the Humbug Scrub area. particularly 
in the vicinity of the South Para River, are exeeMent, and much of die district, 
situated as it is within easy distance of .Adelaide, should prove a most fruitful 
subjea for detailed held mapping and pelroloiricai studies, Numerous other 
Strut] areas ot Archaean schists associated vvith granitic gneisses — ihe Barossiati 

Series-- are to be found scattered throughout the Mount Lofty Ranges, &to it at 
Houghton, F&rtf&ft, Ablate, Yankahlla ( Benson 1909 ) , Meadows Valley 
(Mavcsun 1023), Normanyilte. Mypoirra Tiers, Mount Compaq Mount Craw- 
tor district (England 1935), Tanunda district (Ilossiekl 1925). Ccn^ni (1909) 
has noted a marked similarity in. and certain characteristics pj the r "ij;neoUs*' 
rocks of ninny of these widely scattered areas, which moved liifrti tb posiu'ale a 
jtfwt^e potrovraphic province in which one ma^nia — Hie Houghton niagina--was 
responsible for these gneisses by injection into the r.aros.s : an Schist?. Iu The 
fijvfii of modern ideas on grarritisation. the whole subject of the Tlouffhron uiagffia 
mbdu well he re-exanrncd. 

Tbe present, writer wa;- rrrondv privileged io make a brief \diit of inspection 
of the Little, florae area, about .H tmHes youth of Normnnville. Here metamor- 
phosed ilniemtic ^riis and conglomerates, presumably basal beds of the Adelaide 
Series, are found interlaced and injected or replaced lit-par-lit fej pink pegmatite 
felspar stringers, Immediately east of these outcrops the rocks srrade to phyllitic 
schists with ilmcnite- and masjneiite-bcaring rjmirlz- felspar veirTets and in places 
containing porphyrobkwie pink felspar and bine opalescent quartz. Xearby the 
rocks develop a definite gran i lie ^nei.^sic si nurture and hybrid igneous-looking 
ri'ik types arc developed, c,rj. t broadly gnefssic ''epidotc syenite." The wriler was 
left whh tbe impression that this district should prove Highly instructive for the 
siudy of ;be progressive metamorphism and e.ranitisa:ion of a series of .sedi- 
mentary rocks. 

One oi the best known regions of Archneozoic gneisses in South Australia 
is hi southern and western Kvre Peninsula. Tilley (1921 b) has recognised and 
described four groups or divisions <A rocks from this area which, in descending- 
order of nnHejuity. are: (i) 'flic Hutchison Series, fin The. Flinders Series,, 
(hi! The Warrow Series, and (iv) Th? Dutton Series; 

The Hutchhon Series is considered to be predonrnantly of sedimentary 
Cvtieiil — dolomites, calc-maguerian silicate rocks, paraernei?scs and graphite rocks 
('Tilicy 1920, 1921 ad — and has heen invaded at:d metamorphosed by the granites 
and gnris^ of the Flinders Series, the most widc-soaccd member of 1be {*rt)lip> 
The Warrow Series comprises metamorphosed sediments and is distinguished 
from the Hulchison Series by predominance of massive qaartzite.s iu the former. 
A laler Ih-parTit injection info the Warrow sclvsis of yranite characterised by 
an nbmvlance of tourmaline in its acid differentiates, ro^hed in development ot 
the Duilon Series injection gneisses, 

Tillcy ( 192J b) has described the rocks of the Flinders Series hi detail 
— hornblende and bioiite e/nun'te irneisscs. charuockific bvpersthene and dicpsidc 
Granites, wirh UpKte* and pegmatites, the latter bcinq; eM^rially characterised by 
hornbhnde, Tie concludes that (he jmeissie structure in tbe granites is a primary 
srruerurr roprcscntins a foliation imparted to tin* inrks during consolidation* 
It is siomilieant that the genera! trend of foliation in the Flinders Kan^rc gneisses 
f north-south, with vertical fch sleep westerly dips) is similar lo llie general strike 
ard dip of tbe Hutchison Series sedimentp, which are apparently intruded and 
enrlni-el bv the granite srnciFses at ce"lnin localities in Ihc SleaTord Itay area. 
Tpfey has pnid considerable attention to intercalated bands of basic rock;: in the 
;irid gneisses. r/.?., amphihohtes, hornb'ende gphl^tft pyroxene granuhtcs and the 
like, v/liic!i he concludes to represent metamot phoned remnants of pre-existing- 
basic LfitCt'jUft riK-k^ v, of ^H^htly hiedter aeifliry th;m d.e nono;d gabburd or 
d,nleritic types of intrusive" (1921 b, p. 11), product* of an earlier ron^oliration 
wbieb lta\ ri been etigltlled by the later nci.l ;/rei_-ises. Tie >.n< de=cribed and illus- 
trated leniieidiu- drawn out and comoiu\d bodies of such basic jriGHiSiOU*. wdiosc 
folding is cxphdued as plasuc deTonnation following lii-par-!:t injeclinu and 


magma flow, tJ\ t ptygmatic folding consequent upon intrusion by the granitic: 

it wonld appear thai a good case for granitisation, £g tJ production of Flinders 
granitic gneisses and migmantes by replacement o\ the original ( ? Hutchison) 
Scries, could be presented. The presence of the abundant remnants of basic 
"igneous" rocks— rare components of the nonual Hutchison Series — as xenohths 
in the migmatites suggests once again the possibility that these anrphibobtes may 
represent relict* of the fixation product* of a basic "front" which preceded the 
onset of regional graiuiisation in the area. 

Further studies of the geology of Southern Eyre Peninsula ^rc 
apparently yet required to confirm the stratigra.phic position of the Warrw 
Series and discover the genetic relation, if any. "between the Flinders and the 
Dnttori Series of gneisses. Another region of grannie, rocks KUely lo yield 
interesting results from a detailed invesiigtiion is In eastern Eyre Peninsula, 
in the belt nmuing northward from Port Lincoln to Iron Knob and beyond. 

Finally, brief mention may be made of the granitic ruck* of the Flinders 
Rrnges. Probably the most interesting occurrences yet recorded are those in tils 
central igneous and mctamorpbic complex of the Mount Painter region. Mawson 
(192Jb) has correlated these with rhe Archacozoic Erd, but according to re.enl 
investigations described by IL C Sprigg (1945). in au as yet unpublished report. 
there is here evidence of a Lower FV.aeozoie nge for the igneous activity, possibly 
related to Ordovician orogenic movements.' n Two types of granite b:ue, been 
recognised- — a typically stressed or sheared red granite iu which -sill phases of 
assimilation and gratiitisatiou replacement of the adjacent Proierozoic (Adelaide 
Scries) metamorphosed sediments have been recognised* and a white or leueo- 
granite also intrusive inlo the Proterozoic sediments bnt thought to be younger 
than the red granites. The white granite outcrops arc typically massive, 
unstressed and frequently closely associated with the Thick Quart?! it horizon 
of the series, which is commonly found as xenohths in rhe granite. 

Scattered outcrops of leuerocratic granite arc also found in the Flm.lcr^ 
Krmgcs outside the Mount Fainter area proper. Petrographic details of some of 
these occurrences near Uuiberatana have recently been described by Mawson and 
Dallwitz (1°4S). who consider thchc. inirudons to bfi "in the nature of cupola 
summits above rhe general pluronitc muss of a large scale granitic intrusion'* 
(p. 48) into the thick ProterozoJc and Cumbrian sediments, 

In the Mount Painter area there are numerous good examples oi gradmioi-al 
contacts between gncissie red granite and adjacent quartzitcs and quartzitc 
inctusions in which the strike and dip of gneis; : .osiL_y in the granite and of bedding 
-structures hi the quarlzires remain sensibly constant, and grade imperceptibly I \ 
into the other. Pock exposures in the Mount Painter area are excellent, and, 
despite the relative iuacccssibiiiry and rugged nature of the country, parts <.>:" the 
area would provide wonderful opportunities tor detailed held studies and funda- 
mental research into the mechanism ol emplacement and pedogenesis ot ikv-e 

In the preceding note.s the wri-er has commented on a few areas of Pre- 
Cambrian granites, gi anile gneisses and migmatites which have come to his notice 
and which, m the light of modem ideas, he considers would well repay further 
intensive studies. No doubt mrmy South Australian geologists more familiar with 
the State ihau be, can recall oilier areas equally suitable. It i- to be hoped that 

C*J A tret-jit (leimmration of tin ka'.l/nrnniiui] ratio, on saiiiar4;ite from Mount 
Pu ; uiot (Kleeroan, A. VV.. l<MO, Trans. Hoy. Sue. S. AusL, ?0, J75 177) support* &!£«■*- 
tiun e' a Lower l^sjaeojstfiu ayjft 


they, their associates and stiirlctnts will combine to foster a spirit of more search- 
ing enquiry into this subject of grunirisation and die origin oi Lite granite 


The general problem of tlic genesis of ore deposits raid of the ore-forming 
fluid, and more particularly the origin of those ore deposit? must closely assoeiale.d 
a\ ith granite or with rocks of tin: "phi tonic association/' is still very much a con- 
troversial subject. It is a subject very closely allied to ibe grarrte problem and 
naturally theories of gtauitisatkm and replacement metamorphism are of special 
intercut to the eeonomic geologist. For those who have not questioned the ottho- 
magmatic nature of granite a reasonably satisfactory explanation of the genesis 
of certain o[ the metallic ores typically associated with granite, ct/*, tin. tungsten, 
tantalum, molybdenum, bismuth, fete., has been provided by theories of dilferentia- 
tiou from granite magma, and expulsion of ore bearing solutions together with 
end phase volatiles. and subsequent deposition under resfictcd physico-chemical 
conduioiis. On the oilier hand, if a granite can be formed from pre-existing 
sediniM'ts by the action of suitable emanations, then the primary source of any 
metallic elements which may he fotind associated with such granite becomes more 
obscure than ever. Under these circumstances they can represent either ma:crial 
which has migrated forward and out from the original sediment during gmnitLsa- 
lion* i.e., possibly forming an advancing metalliferous front, or ihey may he 
original components of the granitisaaou emanations. Iu the former case the 
apparent restriction of certain metals, r.*/., tin, io a gran le association becomes 
more pruzling than ever, since other meta's, eg., platinum rind chromium, arc 
found L\ association only with basic magma, 

Kastall lias pointed out another very pulling feature of ine.tyllogeiiesis asso* 
dated with granites, namely, the sponuFc distribution of certain rtiuemls in lime 
and place. For examole. the tiii-tuugs'cn-lourniatine association with granite i3 
fairly common but is found in Western Flurope only with the Hercynian granites, 
in Malaya with late Mcsozuic probably Cretaceous granites, and in Bolivia pro- 
bably Tertiary (f&rgtaU IS4S, p. 27 i. In Western Aivtiraba this association is 
found iu the I're-Cnmbnan granites. Discussions such as these may at first appear 
largely academic, but when consideration is given to the subject from the point of 
view of developing a working basis lor scientific prospecting the economic urgency 
of the whole problem becomes obvious. 

A specific Cast! which immediately springs Id mind is that of the search for 
new ore deposits in the goldfirlds of Western Australia. Potentially auriferous, 
the "kindly" country of prospectors is generally accepted tti he the "greenstone" 
country of the ''islands" mentioned abov*. The surrounding granitic and gncissic 
country, on the other hand, is usually eou-adcrcd to be non-auriferous, despite the 
frequcnt presence of quanz n-efs whhhi L. Accenting the principle of the forma- 
tion of the ;4rnmiic bodies by replacement of pre-existing rocks, there arises the 
possibiliiy thai pre-existing gold also occurred within ihese -oeks and much of the 
ore found in known auriferous belts in the greenstone areas may represcin 
''throvcouts"' formed by the forward migration o ; go!d from the granki^ed nreas. 

It is conceivable under such circumstances thru remnant of such original 
gold-bearing rocks and local concern ratums of aurifernus material may have heen 
cammed back or otherwise retarded from lvugniiirg mil of the: area of granitisation 
send thus may occur withiu die granite areas, piivticulark within the marginal 
gueissic nrigmatite zones- The ore bodie,s of the: Kdna May { W.A. J Amalgamated 
Hold Mines, at Wcstonia in the Vilgarn GoldfJeld, containing gold, tungsten and 
iiirHybdiluun, arc found in biotile gneiss country. !n a number of other scattered 


localities Eli the northern Yhgarn Goldfield small gold workings arc to be found in 
wbat are considered to Be relict greenstone lenses either partially or completely 
enclosed by granitic gneiss (Maiheson J 940). 

The question of the genesis of iron tire deposits has been a subject tor active, 
controversy for many years, in an investigation of iron ores assoeia:e 1 with 
banded hunatiie quartzites or '"jnspiir bar/' in Western Australia srveral years 
ago. titc wrirer noted evidence suggesting at least two generations oi iron con- 
Centr.aHoii : (a) a molccidar repIaceraenL of silica bauds in original magnetite or 
hematite quart/dies By granular crystalline hematite., arid (b) later uuplaccmetH 
of discordant ieiVdetdar bodies or veins oi coarse crystalline specular or micaceous 
hemacke often intergrown with quartz. At the time of writing this replacement 
was considered to bo a supergene process, though possibly assisted by heated 
magmadc waters (Miles 1941, p. 197). The iron ores ot Koulyunobixing, 35 miles 
north-east pi Southern Cros-. hi the Vilgarn (jolddcld. show these features in no 
small decree. 'Die Koclyanobbing Range forms a long narrow belt of vertical 
dipping' banded iron formations in greenstone country fringed both on the east 
and west by broad areas of granite and/or grists. There is reason to believe that 
large areas of ferruginous rocks — basic schists, lavas and banded iron fcrina- 
lions — must have been replaced during the emplacement of all fbis granitic rock. 
The writer ha=. seen in o'her parts of die district excellent examples of gsennUsttil 
0* denatured banded h-on formations in which ail the iron ore has been removed, 
probably as sombre, ha.idcs, leaving either a bleached whke quartzite or in some 
cases banded Huonte quaruue (Miles 1<M5). It is possible diat the iron migrat- 
ing forward wall the progressive granit'tsation of country rock at Koolyamdddag 
has foitned an advancing "terric front'' which rrwv have been responsible for 1h-- 
coarse quartz-specular hematite bodies, ami possibly even for the carrier main 
gnrnular hemadt/- replacement bodies mciuiond abo\e. The tendency for ali this 
iron to coneenuate within a restricted zone, i.e., the. original banded Wnvuke 
qtiamite horizon, may p^.'htb'y be explained partly by a chemical affinity uf the 
migrating material for the a ready highly ferruginous handed sediment, and partly 
hy the structural Control (damming a:. an) exerted by these banded ferrngh-ais 
quart?:! re beds. 

This subject of lh? rc'ad^vhrp jit any) oi adjacm: granite bodies to Prc- 
Cnmbnan handed iron ore deposit:, iucfi as as. Yampi boon::. Kuclyanokolng, 
\VtlgpJ Mia. Trdierhig Uangc in \\ v.Tern ;.;■> raba and In the JVI:dJki;cak !hm;w. 
h''«ud. Acs.ndia, should h^ vvb v.yir-Ui Yn\'&J»\' invwrgad n\i m 

( '• n -i'.h-rahon of dav g.anhc problem U a mo>f ah*rtrt£t<jfj occupatina .Y - l.odi 
da? rVfOFitcii stiatEsci: an 1 . *1v economic r r -i'"dc :;i-i\ This coiimry provide, sc-v'h* t:;o-:nou:; amotmd of b:temive inveshgadon of An: granites, bur partuuk .y 

il:e! rlaa cranh^ nivl d:Hr a:: o^hnes. ik>Ci ihdd ;md thaaiicj laho-a^'^ 
^uvlles r-;r la-nirch The wri'er wo:thi pavticiihirly :-.'•;.., Qi u ne:e.ii[y h=r red 
wod; whh c:nv c ut rca-rdrng of fac^ foia p.r^garQjicta tiff a.var;;tv g.u.o r'cal ;;:ay.= 
o: ; h^:h reylonrl and dekiilpd ^ca'es us a preiimmury to chvnrcal and penological 
studies, a prime rcq;:h-::me::: which 1 a-- r.oi ahsays f;:dy appr-ciai^'d by some 
u-a'.'t ioic worh^rz. iu carrving oat rids ty})?. of mvestief.iLa-.u careful and a^rurm 1 . 
ilvcrv':t'eu of la:'S is imt rc-jcire-h fohowrd ly ;::» aupai dal {U^^man'-M id 
these farrs. togahc'- Xvkai a i and logical disv^-oion rd the impdeadoiis 
and deductio!'S lo he drawn from th^m. in ihfs connect'on i; may he rfojl oui r -i 
place to dran T attej:ti.^i to ti^e criiieal review <if same r-cctn works r,n\ the stig- 
gested specifications for a sai^factorv report which were i^ucd a few vrrr>; f - 
bv F. RGrot^ (1911 J. 


tty means of researches such as ihcsc we hi Australia should be able to con- 
tribute notabiy to world Jctu^JeiJg^ (Sn :his most fascinating and contentious of 
questions. "What is granite ? M — and at the same, lime take a -worthy place with 
other observers in countries in which the grange rocks. especially thore of the 
Archaean shield, arc abundantly dismayed. 

The ideas discussed in the preceding pages are not new, as has been clearly 
indicated, nor are they prevail to be ot general application. M'hii.= t some of the 
suggestions put forward by the present writer are frankly admitted to be pure 
speculation, not based on any personal experience. However, it any Of these 
suggestions are. sufficient 1 }' arresting to simulate amongst geological workers, 
particularly those in South Australia and Western Australia, a renewed nucrest 
in, and a fuller awareness of, flip prub'ems of the granite of both Pre- Cambrian 
and later ages, he will feel amply rewarded, 


The writer is Indebted to Mr. S. IF Dickinson and .)r>. L. K. Ward and 
R. T, Prider for constructive criticism and many helpful suggestions during 
revis'ou of the text of this paper, 


Alderman", A. R, 1938 Augen Gneisses in the Humbug Scrub Area. South Aus- 
tralia. Trcns. Roy, Soc. S. Aust., 62, 163-181 

IUcklund, 11. 193S Zur lt Granitisat:onstheo-ie" GroF Fdren Forhar.. 11, . 60, 

Rexsox, W. N. 1909 Petrographical Notes on Certain F re-Cambrian Rocks of 
the Mount Eoflv Ranges, with special reference ro the Gcologv ot the 
Houghton District. Trans. Roy. Soc. S. Aust., 3ft 101-140 

Duxk, F A. 1942 Cranke and M agnation and Mctamorphism. Eeon. Geol.. 

Rlljs, IF A. 1939 The Geology of the Yilgarn Gold-held, South ot the Great 
Eastern Railway. Geol. Surv. W- Ausi., Full. 97 

ExolA^d, IF N. 1933 Petrographic Notes on Intrusions of the Houghton 
Magma in the Mount I oily Ranges. T-ans. Roy. Soc, S, Au>F, 59, 1-15 

Grun.T, F. F. 1941 Fomunion of [^ucous-lookiu^ Rocks by Metasomatism: 

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Roy. Soc. S. A nst., 49, 191-197 
KrxNrnY. W. O., and ANnr:*sox T , E- M. 1938 OwS&U Payers and the Origin 

ot Magmas. Hull. Yolc. r Series IF Tome III, 24 82 
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Mawsok. D. 1923a Notes on the Geological Features o' the Meadows Valley. 

Trans. Roy. Soc, S. AusU 47, 371-375 

Maweox, D. 1923b Igneous Rocks of the Mount Painter Belt. Trans. Rov, 
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Ma\vso\, CK, and Dallavitz, \y. |>, 1Q45 The Soda-Rich Leucogranitc Cupo- 
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Mills, K. R. 1941 Magnetite-Hematite Relations in the Banded Iron Forma- 
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Mjlls, K. R t 1946 Metamorphism of the Tamper Bars of Western Australia. 
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Prtdek. R_ T. 1941 The Contact between the Granitic Rocks and the Cardup 
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Prtoef-:, R. T. 1945b Chartiockitic and Related Cordienre-hearing Rocks from 
Dangin, Western Australia. Geol. Mag., 82, 145-172 

Rastail, R. II. 1945 The Granite Problem. Osfi. Mag., 82, 19-30 

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REAn, PI. H. 1944 Meditations on Granite. P.\ 11. Proc Geol, Asso., 55, 


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Acad., 48b. 231 

Rkvn'oius, D. P. 1944 The South- wcsLcni End of the Xewry Igneous Com- 
plex. A contribution towards the Petrngenesis of the Granodiorites. 
Quart. Journ. Geol. Soc, 09, 205-240 

Rrvvoi.os. D. L. 1946 The Sequence of Geochemieal Change- leading to 
Granilisatton. Quart. Journ. Geol. Soc, 102, 398-438 

Spriog. R. C. 1945 Reconnaissance Geological Survey of 1he Mount Painter 
Area. [ Unpublished ) 

Tilllv. C. E. 1920 The Metamorphism of the Pre-Cambrian Dolomites of 
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TtLLF-V, C. II 1921b The Granite Gneisses of Southern Eyre Peninsula (South 
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77, 75-131 

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Svecofenniden in Finland 1, II. Bull. Comm. Geol Finlande. Xo. 89 

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Wf:eLM'--\.\, C. E. 1935 Caledonian Orogeuv in Christian X's Land. Medd, 
om Gronl., Bd. 103, Xo. 3 

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Medd. om Gronl,, Rd. 113, No. 2 

rans. Knv. Soc. S. AuM.. VM7 

Vol- 71. Piatt l 

Air p|iotQ o Granite Area, ;m >t_a i 7 mile* north of M6UI1I Painter, Flirulers Rangx*. 
North at top of photo. Shows leueo^raiiite ( lower right half of photo) intruding 
.oi<i replacing portion of the 'Thick Quartzite horizon (upper left). Abundant 
pegmatite off-shoots are "advancing 1 ' ahead erf the granite. Numerous oriented 
remnant- nt sediment c;oi he *ceu enclosed in granite (lower left). Soulh-suinli- 
easlt-rlv unvoted dark I lilts in maoite reprL'scnt major jointing plane--. 

R,A>A.F, Photo fVerticnll No. H)2, Run i37, L»t. .>o<)7'S.. Lon#. Hft^JffE- 
}.Vr MY R, li April 1545. Scale approximately 27-5 chains to the inch, 


By J. A. Prescottand C. E. Lane Poole 


Apart from its native grasslands and forests, the continent of Australia originally offered so little in 
the way of plants of economic value to man that plant introduction has always been of great 
importance in the development of the country. The botanic gardens, early established in the capital 
cities of the colonies and at Darwin, were originally essentially acclimatisation centres and testing 
grounds for introduced species of plants, although they have generally lost their importance in this 
regard and this function has been replaced by the plant introduction activities of the several State 
Departments of Agriculture and the Council for Scientific and Industrial Research. 



By J. A. Prescott< 3 > and C. R Lax*: Poole <-> 

[Read 14 November 1946] 

C o n i Exrs 



Piims ■>'('.(! ktia 

Punts' ru'iarifuxis 

P-nns pvnnstcr . . . . . . . . . . . . . . 

Punts nifjra 

Phnts haleprns's 



Aesthetic p'ant ngs . . . . . , , , ► » 

Forestry p'antatious .« . . . . . . . . 


The growth at Monterey Pine in relation to Environmental 
The Choice of Appropriate Species for Soulheni Australia 
AcKKOWLKii IMf-.VJ'S - -* 

R.EFliRKNCES . . ,,*. . . . . . • 

Solaces of I x folimatiox 





.. 70 

.. 72 

. . 74 

. . 77 

.. 78 

.. 73 

.. 79 

.. 79 

... SI 

., 85 

.. 85 

.. 87 

. 89 

. . 8'.) 

.. 89 

Apr-jt from its native, grasslands and forests, the continent of Australia 
originally offered so little in the way of plants of economic value to man that 
plant introduction has always been of great importance in the development of the 
country. The botanic gardens, early established in t\ie capital cities of the colonies 
and at Darwin, were originally essentially acclimatisation centres and testing 
grounds for introduced species of plants, although they have generally lost their 
importance in this regard and this function has been replaced by the plant introduc- 
tion activities of the several State Departments of Agriculture and the Council 
for Scientific and Industrial Research. 

0) Director, Waite Agricultural Research Institute, 

V§ Formerly Inspector-General oi Forests and Acting Principal Australian Forestry 

Trans. Roy. Sec, S. Aliitti 71 (I), 25 July 191! 


111 the earlv years of coloration \h%t& niu.t have been extensive trial and 
error hi these activities, but the exigencies ot" quarantine and the exhaustion of 
the more obvious introductions have made it increasingly necessary to place plant 
introduction on a more logical footing. Some highly desirable introduction^ such 
as the sov benn ftavC proved exceedingly difficult, while there is no doubt that 
others have failed through a lack of knowledge of soil and climatic requirements, 

In a recent discussion on ecology and the study of climate reported in Mature 
by Day- !i) (1946) emphasis was paced on the need for the growth-cycles of 
introduced spedes to be in phase with the annual seasonal cycle. Two examples 
are given of such lack of harmony in Britain. The European larch begins its 
growth nmcli loo early, whi'e the Corsican pine continues growth until much too 
late. Because of this they suffer seriously from fro-t in many situations. This 
tends markedly to restrict the areas within vchieh they can be grown successfully. 

It J* th.e pmpL'se of Ihe prescu: eontrbuHon to rcv ; ew the climatic require- 
ments of a number of species of the genus Pinus which thrive in the Mediter- 
ranean region or in region- having- a similar climate such as California, and to 
determine "how far such equates art- reproduced in southern Australia- In the 
light of actual experience witli these species in Australia, the analysis may reveal 
some genera! p-rinc'ptes which may be oi service in such plant introduction work, 
The species chosen are: 

Finns nitl'iaiti D. Don, die ilomertw pine, from California. 

rftitts eurtcri^nsis C. Smith, the Canary Island pine. 

Finnjs pinaster Aiton, the cluster pine, of southern France, Portugal and 

Finns i : ;i:ra Arnold or* Pinus l^ricio h'o:;\:L rite blacker Corsican pine, also 

of the Mediterranean region. 
Pi*'iis linlefcnsis Miller, the Aleppo pint- of the eastern Mediterranean. 

The Monterey and Canary Island pints have very restricted native habhais. 
so that the climatic conditions of these environments can be very narrowly defined* 
The other p : ues are more widely distribute,;, and the associated climates are mere- 
fore nvi quite so easily determined. 


in dculim; v/hh perennial plants, uwf&rtitarvis and moisture eondhi-v. > 
throughout the'year are ot Importance. Tolerance to low or hi:*b temperatures 
mac be important, and greater empnasis has therefore been placed in this sdidy 
on. "temperature. Isaiufa i h- not, umrtwer. completely overlooked. 

In dealing with temperatures, m*r- lias been made of the convenient method ot 
wave f^rm aaalv^is adc-pred by 1'r ckcuu (UC2). which enables \h# twelve mean 
monthlv temperaturr:; to be reduced to ihe ilim* values of annual mean, amplitude 

Map? ot L'.ic M;''.1Hcna"Ci;U rrcden, .shcmir<_' 
(a) Natural dktribution tri f4*MJ ^eHtfr** /\ f^mtca^ K ifystft < r: - /:Or/v;;^. 
(10 Mean amiunl tempera lure. 
ic) Mean unntia* atnplitud^ of tvicpc^atJire. 
i<l) Teinperar:tre ehr^o &c)nrtiS&prf in turns til 1 ;i ^ hi ikiys tehind jfohf r«diatiuu 

' "i' Sen aho t)aj (ls-5). 


ff*J £ car ar iff ?>i$ . .._ / j 

I'M ; - ■ ■ 

Fig, J 


Fig-. 2 

Map oi the Mediterranean region, showing mean animal rainfall in inches 
ba*&[! on data of Kencirew tl'.WJ, Brooks (1932), and Tlcrbcrtson (P>01). 

and phase. It is assume.! that there is some correlation beiweeu daily amplitudes 
and HTiuuai amplitudes. 

Of importance in the case of nioisutre conditions arc the total amount of 
effective or influential rainfall and Its distribution in time. Ln the Mediterranean 
climates under consideration, the rainfall ha- a marked vvinter incidence and the 
igftgth of the rainy season becomes au important index. To secure identical 
combinations of all these factors is fiat always an easy matter, but it is essential 
to secure identical combinations 01 at feasl mean annual temperature and ampli- 
tude with the appropriate seasonal rainfall. One difficulty is to assess the con- 
tribution of cloud and fog to moisture supply. This h particularly so in the ease 
(if the Canary Island pine and the MoiHerey pine, i: is probable that in these 
cases, in the Australian environment, somewhat higher rainfall should be allowed 
than is recorded for the native habitats. 

Maps 61 the Mediterranean region, illustrating the geographical distribution 
of four of these pines, together with temperature characteristics and annual rain- 
fall. a"e given in fie, 1 and 2. 

J-'inus raujaxa ( Monterey pine) 
Phius rmliufa is a native of California and shares with Cuprcssus macracarpa 
the verv re>lricted habitat of the Monterey peninsula. Being essentially a coastal 
species occurring- near sea level the temperature conditions can be readily ascer- 
tained without interpolation, and the rainfall conditions arc similarly reasonably 
well known. The other habitats of this pine are Ano Nuevo Point, San Simeon 
Bay (Cambria), two of the Santa 'Barbara Islands and die Island of Guadalupe 
off the coasl of Southern California. The climatic conditions on these islands 
have r.ot been ascertained, hut they are likely to include low amplitudes and a late 
phase. Attention has necessarily been restricted to the occurrences on the 

In Table I are given the essential data for Californian localities near the zone 
of natural occurrence of this pine. Santa Barbara and San Luis Obispo have late 
phases associated presumably wiih greater oceanic influences. All !he amplitudes 
arc quite !ow. The annual rainfall on the Monterey peninsula has heen estimated 
by Mason (19.34) to range from 18*8 inches at Carmel to 25*7 inches on Huckle- 
berry Hill, with almost daily summer fogs. 



Temperature ami Rainfall Characteristics cf Lowhies [n o\- 
near the Caliiornian Zone of Occurrence of Pirns racUoia. 

Temperature Charac 



all + 


T1 stlC> 

Phase, l£g 

bell r.d >-,-mV 

l^r:gth 0) 

Annus! Alt 







of season 



a F. 


Ifi *, 



San Luis Obispo 

.... 58-5 






Saittn Barbara 

.... 59*8 






Santa Cruz ... 

..,. 57-3 






Del Monte ■ 







For practical purposes, therefore, may be sought zones in southern Australia 
having the following temperature characteristics: 

Mean annual temperature .. - — 56-60 F. 

Amplitude <• • ---- » 5'5-7'G F. 

Phase 35-50 day? 

Parallel zones (fig. 3 and 4) are found to occur in the south-west of Western 
Australia arid in the "coastal regions of South Australia, Victoria and northern 
Tasmania. Kangaroo Island and the islands in Bass Strait aUo come within the 
zone. The climate is thus essentially maritime. Rainfall conditions within this 
zone are, however, much more favourable than in most of the Caliiornian 
localities, and it may be assumed that adjacent zone-} with annual rainfalls of 
25 to 30 inches may come within the favourable zone, providing soil conditions are 
suitable, it is known from Australian e\periencc that this species is rather exact- 
ing in its soil requirements, atld throughout this discussion it must be bom- in 
mind that eclaphic factors are deliberately excluded from consideration. Il isof. 
interest to list Australian stations winch oiler parallels to these Calu'onran 

Table 2 

Temperature end Rainfall Giaraeteristii.^ of Australian Local itk'S having 
Temperature Regimes similar to those -of Caliiornian Stations in or near the 

Habitat of Pirns r&ii(.'fa. 

Mean annual Alton nrnm.*:! Lmyth oi* 

lemiieruture Amplitude "pllgSfi rainfall wet seu>oji 

X.oc;.1tty ° F. a K. d*ys ins. month;. 

Kdcl>*tonc Point, Tas. .... 56-6 6-3 42 29-4 12-0 

CfrrrJt, King Island, Tas. - , S5*7 5 ; 3 47 34-1 12-0 

Mbanv. W. Aust 59-9 6-5 45 W~2 1J-4 

Eclipse Island, \\\ Aust SSKS S-fi 52 32-7 12-0 

Karridale, W. Aust. 59-8 6-6 $4 47-8 9-8 

Cape Borda, 5. Aust 58-6 6-6 41 24*8 7v9 

Kingscote, S. Aust - .. 5*M M 41 I'M b-8 

Cape Northumberland, S. Aust 5(r9 n>3 38 2f>i> 1.0*6 

Robe, S. Ayst !.. ... 57-9 6*5 3? 24*7 0-3 

Mount Burr Forest. S. Aust , =S-5 5-7 36 30-7 9-4 

<") The mean length of season in this and the following- tables is based oi; mean 
monthly values for rainfall and probable evaporation. It would be preferable to use the 
mean of the length? of individual seasons, but this reformation was available in or.Iy a 
few cases. The estimates are likely to be somewhat loo high, particularly for the higher 


it will be seen that the South Aus- 
tralian stations approach rno=t closely 
the Caltfornian ones in respect both of 
temperature and rainfall. The close 
parallel between Kingscote, Kangaroo 
Iblaud and Santa Barbara may be 
noted as one example.. The Australian 
stations have also longer rainfall 
seasons and a greater certainly of 
summer rainfall. This may well com- 
pensate for the absence of summer 
fftgs in the Australian environment. 


(Canary Island pine.) 

This pine, like the Monterey pine, 
occurs in a rather restricted zone, but 
as aUimdc plays a part in the de- 
termination of its habitat, it has proved 
necessary to interpolate some of the 
climatic information available in the 
standard textbook.? of climatology and 
the report of F>\1es (1932). 1 1 lias 
been assumed that this pine finds its 
optimum temperature conditions al 
altitudes between 2.000 feet and 6,000 
feet, with 4.000 feet as a probable 
ideal. Although the coastal stations 
of tfe Canary Islands have a mari- 
time climate similar to that of coastal 
California, there appears to be a sig- 
nificant increase in the annual tem- 
perature range with increasing height 
above sea level. No part of Australia, 
shows the -harp changes in topography 
charrxtm'stic of the Canary islands. 
Lyie.s assumes a probable annual rain- 
fall of between 12 and 30 inches. The 
len^li of I he rainy season cannot he 
predse'y determined from the avail- 
able data, but it appears to be in the 
neighbourhood of eight months, with 
a create r certainty than in Australia 
1 1 f ul^cnce o i rain in the summer 
months. The temperature data which 
are appropriate to rhe discussion arc! 
eiven ?s Tabic 3. 












x^ 8 *^,*^^* 

Fig. 3 

Maps of tile south-west part of Australia, 
illustrating localities where temperature con- 
ditions are similar vytth respect to mean, 
amplitude and phase to those of the zones 
of native occurrence of P. radial a and 
P. citr.ariaisis and the Portuguese habitat ot 
P. fniuufe}-. The numbered lines are isohyets 
of rainfall in inches. The line of circles 
parallels temperature conditions with respect 
to appropriate combinations °f mean and 
amplitude in ihe ease of P- caiuit'it'jisis. 


Table 3 

Tempe-ature Characteristics in the Neighbourhood of the Zone of native Occurrence oi 

Pinks C'Ti'oricnsis, 

Santa Cruz. Tenerifte - , 

Puerto de Orotova, Teneriffe 
Guimar, TcnerilTe 
< : ') Vibiloi-, TenerifTe 

Las Canadas, Teneriffe 
Las Paluias, Grand Canary 

Funchal, Madeira 

Agaclir, Morocco 
Mogodor, Morocco ,... 

From the data available it may be assumed drat the temperature requirements 
of Finns canan'pnsis approximate to the following conditions on TeneriiTe : 

At 2,000 feet 
At 4,000 feel 
At 6,000 feet 

SUfl level 

Annual meat: 
a F. 

c F. 


lag beinpfl 

solar radiation 

















11 -6 






























Fig, 4 

Map of south-eastern Australia, illustrating localities where 
temperature conditions are similar with respect to mean, 
amplitude and phase to those prevailing in the California 
home of P. radiaUh The areas are shaded and include 
Kangaroo Islam!, the islands of Bass Strait, certain coastal 
regions of South Australia and Victoria and the north-east 
corner of Tasmania. The numbered lines are the isobyets 
with annual rainfall expressed ui inches. 

It is possible to 
obtain appropriate 
combinations of mean 
annual temperature 
and amplitude over 
the Avhole range., and 
these combinations 
have been plotted on 
the map of Australia 
and are illustraled in 
the maps of tig. 3 
and 5. The nearest 
approach to similar 
conditions occurs near 
the coast at the 
extreme western and 
eastern geographical 
limits in Australia. 
Stirling West iu South 
Australia. Terang in 
Victoria, and haun- 
eeston in Tasmania, 
closely approach in 
some respects tin: 
mean climatic habitat. 
Some selected values 
of this kind are given 
iii Table 4- 

& Calculated from partly interpolated data. 




behmtl solar 

1 radiation 


Length of 

wet season 















23 -8 


5 f M 



37 -.3 

















Taw,e 4 

Temperature and Rainfall Characteristics of Australian Localities showing 
some Features similar to those of the Zone of native Occurrence of the 

Canary Island Pine. 


Cape Naturalise, \\\ Aust. 

Busselton, W. Aust 

Bombala. N.S.W. 

Gabo Island. Vict. 

Latin eeston, Tas. , 

Tterangi Vkt - 

Stirling West, S. Aust. 

In rflogt of these cases rainfall may will be excessive and more favourable 
conditions may possibly be found in somewhat drier zones adjacent to the appro- 
priate temperature zone. 

Pinus i-ixaster (Cluster pine) 

This pine occurs naturally and is cultivated success Fully on the Atlantic 
coasts of southern France and Portugal. Certain strains are characteristic of 
Mediterranean France. Spain. Italy and Cor.sicn. The species also occurs in 
Morocco, in fairly humid environments in the mnunUvns up to a height of 7.000 
feet mainly ht the middle Atlas and m cerlain localities in the 1\if\ It is rare in 
the Grand Atlas. It docs not generally occur east of Italy or Tunisia, but is said 
to occur on the coastal islands of Dalmatia. The African occurrences are not 
regarded as important, hut they afford evidence at least of tbe climatic range of 
the species. On the whole, the cluster pine therefore belongs essentially to the 
Western Mediterrauan. It is regarded as bein^ much less drought resistant than 
the Aleppo pine. Finns hatepcusis, Avliicb is much more widely distributed but 
which does not occur in Portugal, 

On the basis, of its distribution two races are recognised: mttnlinia and 
iiicsorjucnsis. The former race belong to the Atlantic coast, the second to the 
Mediterranean regions and probably also to Morocco and Algeria. The Moroccan 
strain is said to be more tolerant of calcareous soils than are the principal races. 

The temperature and rainfall characteristics of typical localities are given in 
Table 5. The wettest month ill these regions is October; autumn and not winter 
is the wettest season. 

Tablk 5 

Temperature and Rainfall Characteristics of Localities in 
or near the Zunc of native Occurrence of Pinn4 pwwfitt*fi 

Tempera hire 

Atlantic Environment— 
B'o;deiU 1 x, France 
Arcachon, France 
Biariitz, France .... 
Lisbon, Portugal 
Oporto, Portugal 
Coimbra. ForhigaJ 
Santiago, Spain .... 

° If. 


PJinse, lay; 
Inbind snlar 

in c lies 










It -5 



















Ockibti Length of 
rajnlaH rainy sgastBI 







Mediterranean Environment— 

Var, France 

Nice, France 

Toulon, France 

Marseilles, France 

lie cie Levant, France .... 

Montpellier, France 

Cai Corse, Corsica 

Ca^ Fertusnt'o, Corsica..., 

lie-; Sanguinaires, Corsica 

Ajaccio, Corsica , . 

Ra=tcJTea, Corsica 

Corte, Corsica 

Genoa. Italy 

Sessari, Sardinia 

Tn Corsica, the zone of P. pinaster occurs ai ali.itu.lcs of from 1,300 to 
3,500 feet, when it gives place to P, larlclo {P. nigra). It is probable that 
originally /', pinaster extended to sea level. Corte is near the average for the 
Cofsican habitat with an annual rainfall of 34~5 inches, a minimum monthly rain- 
fall of 0-5 inches in July and a maximum monthly rainfall of 5*6 inches in 
November. An estimate of conditions in Morocco, near ll:e north African limit 
of its occurrence, is possible from data available for Dayet Ashlef. Estimates for 
the temperature characteristics of the Corsiean and Moroccan habitats are given 
ill Table 6. It is difficult to predict the correlation between amplitude and phase 
and altitude without some, local data, but the data for Scssari in Sardinia a: an 
altitude of 730 feet is: of some assistance in the case of Corsica. 

Takle 6 

Estimates of Temperature Characteristics of Corsiean and Moroccan 
Habitats of Pwus jifa&tiU'f. 












































12- e 








21 -6 



















59- y 














Sea \p?$ 

fsjaay ranpe 


Mean annual 


6P F. 

55° V. 

51° F. 

S3 B F. 

Amplitude - 


13° F. 

15° F. 

14* F. 

16° F. 



41 days 

35 days 

33 days 

40 days 

The f blowing general limits 'nay be suggested for the temperature chcrac- 
teristies: of the zone of P. pinaster: 

Mean annual temperature - .. .... o4 — 61' F. 

Amplitude .. lO-tS'F. 

rhasc .. ., .- -- 31 39 days 

It is possible further to separate the French and Mediterranean zone from 
the Portuguese zone in order to define the temperature conditions more narrowly. 
For the Mediterranean zone these may be roughly derined by ibe fu'.lmvm* limits 
and their linear interpolations : 

Mean annua! temperature 55* F. 61 J F. 

Amplitude 14 a F. 13' F. 

Phase 30 dux* A\ days 

1'or the Portuguese zone the com di lien Is; can be more narrowly defined : 
Mean annual temperature .... . . 39 a F. 

Amplitude .... 10" F. 

Phase -.- ... .... ... .... 36 day* 

Thesw* latter conditions, with adequate annual rainfall, are closely reproduced 
by conditions at Collie, Bridgetown and Donnybrook in Vvestern Australia, re!a- 


tively near to the coastal plantations of this species of the Slate Forest service. 
There is an approach to these conditions m eastern Gippsland and the adjacent 
coastal regions of New South Wales. 

The relevant data for these Australian stations are given in Table 7, where the 
month ot April corresponds io October in the norLherii hemisphere. 

Tautj; 7 
Climatic Data ror some Australian Stations corresponding in 

Temperature Conditions to Portuguese Hahitat 

M f an 
a K. 




Western Australia — 

Bi klsrefmvn 

Victoria and New South Wales — 
Gabo Island 

Moruya Heads 

The rainfall distri- 
bution in the eastern 
AustiT.l'nn stations 
cannot he said to 
approach ai all ihc 
conditions in Portugal, 
notably in the absence 
of a true drought 
period in summer. 
Parallel conditions are 
therefore restricted to 
Western Australia. 

For the Mediter- 
ranean zones there ii 
v. wider rnn^e of 
parallel temperature 
eonclndons, providing' 
only mean and ampli- 
tude are taken into 
consideration. There 
is an approach to 
equiva'ent phase con- 
ditions in northern 
Victoria and on the 
\wsictu side of the 
Southern tableland of 
_W.v S-.urh Wales, of 
which I'-urriujuck and 
Addons may be laken 
u.-t representative. The 
appropriate combina- 
tions oi mean and 









5 lor P. {'ina,<M\ 

An'i] month 

irifj'tirs inches 



of season 



Map of south-eastern Australia, illustrating localities where 
temperature conditions with respect to mean, amplitude 
and phase arc similar to tho^e prevailing in rfjf habitat of 
P, ca:urruvis:s in the Canary Islands. The areas are shaded 
and include areas in eastern Victoria and the south-east of 
New South Wales. The line or circles indicates paraKd 
conditions with respect to mean and ampltrede only. The 
numbered lines are isohyets with annual rainfall ofcfirfeesttcZ 
in inches. 


amplitude In temperature occur from the north of New England in an arc reach- 
ing as far as Rutherglen in Victoria, 

The conditions at Adelong and Burrinjuck are given helow in Tabic 8. 

Tanus 8 

Climatic Data for two Stations in Nc -w South Wales corresponding 
hi Temperature Conditions to the Mediterranean Habitat of /'. phmsfer. 


Mr an 

TVmf) ciature 
ArmTniile l*hase 



in fall 



of ieason 


... 57-9 

1,5-3 30 






... 58-0 

13 -0 30 

33 -K 




The conditions of evaporation at both these centres i$ such that uear-diought 
conditions prevail during three summer months. H is probable that appropriate, 
conditions occur near these localities at higher elevations. The information con- 
cerning the climatic conditions under discussion are illustrated in fig. 3 and 6, 


(Black pine, Corsican 
pine,, Austrian pine) 

The climatic condi- 
tions under which the 
black pine grows in 
its natural habitats are 
not so easily defined 
owing to the paucity 
of direct meteorologi- 
cal data, associated 
with the fact that 
these habitats are at 
some elevat'on rang- 
ing from approxi- 
mately 600 feet in 
southern 3'Tancc. and 
Australia to 6.000 feet 
in Cyprus and Ana- 
tolia. No climatic data 
have actually been se- 
cured relating - directly 
to one of the known 
habitats of this pine, 
but rainfall require- 
ments are generally 
regarded as high, in 
the neighbourhood of 
50 incites per annum. 

The main centres of 
distribuiron are wide- 
spread but obviously 
localised. They include 
southern France, in- 
cluding the Cevennes 
and Pyrenees ; Corsica, 

Fi£- 6 

Map of south-Pattern Australia, illustrating localities wherc 
temperatnte conditions with respect to mean, amplitude 
and phase are similar to those prevailing in thp. native 
ha'hifats of P. pinaster. These areas are shaded. The area 
marked M is just, north-east of Canberra, and reproduces 
the temperature conditions of the .Mediterranean habitats. 
The area marked P, near Adelaide, reproduces the Portu- 
guese conditions more closely. The line?, of circles 
indicate areas where the Portuguese conditions are 
paralleled with respect only to mean and amplitude. The 
areas enclosed in heavily dotted lines indicate a^eas where 
the Mediterranean conditions are similarly reproduced. 
The numbered lines are isohyeis c-f mean annual rainfall 
expressed in inches.. 


















Sicily and Calabria, Austria and Dalmatia. and the region of the Taurus moun- 
tain s" including Cyprus and localities in Syria. The species also occurs in the 
Crimea and in the Caucasus region and in the Balkans generally- An isolated 
occurrence in the Rif of Morocco at 5.000 feet links Africa with the Spanish 

In order to secure an approximation to the temperature conditions of these 
habitats, correlations were graphically established between altitude, mean tem- 
perature, phase and amplitude. Phase tends to be late both at sea level and at 
very high altitudes, and to be generally correlated with amplitude for geographi- 
cally grouped localities. 

On the basis of these correlations the following estimates of temperature 
conditions appropriate to Finns nigra have been obtained: 

Table 9 

Temperature Characteristics associated with Habitats of P. nigra. 

Mean of all estimates 
Mean for lowest recorded attitudes 
Mean for highest recorded altitudes 
Highest temperature estimate (Cyprus) ,. 
Lowest temperature estimate (Austria) .,. 

The high values for amplitude suggest that Prnits nigra is associated with a 
much more continental climate than the pines that have so far been cousidered- 

No Australian locality has a temperature range as high as that of these 
habitats of Pinus nigra. The nearest approach is in the highlands of New South 
Wales and Victoria. Tasmania is too much within oceanic influences to afford 
parallel conditions. 

Comparison of the above characteristics may be made with the following 
for Australia: 

Mount Buffalo, Vict 

Oraeo, Vict. 

Nimmitabel, K.S.W. 

Crookrveil, N.S-W. 

Wnrataii, Tas 

Pintjs halejpensts (Aleppo pine) 

The Aleppo pine is much more widely distributed than the other pines under 
discussion. P. pumstcr and P. halepaisis occur immediately below the zone of 
P . nigra, the former in the western and the latter in the eastern Mediterranean. 
According to Bean (1925) this is the commonest pine on the south coast of 
Europe, and is at its finest on the Dalmatian coast. It is more drought resistant 
than P. pinaster or P. nigra, and its temperature requirements are continental 
rather than maritime. 

Conditions for two extreme localities may be quoted: 



a p 




















rnin fa 11 









Ragusa (Adriatic coast) 

Bon Taleb (Algeria, 4,100 ft. altitude) 

This latter station has an eflcct : ve rainfall period of approximately nine 
months. Temperatures of typical localities for this species are quoted in Tabic 10. 


° F r 

6 l\ 


S3 -7 









58 -7 

20- 1 









IB- 9 


n F. 














Table 10 
Temperature Characteristics for Habitats of P. hafc/?ensis. 


Mean of coolest conditions 

Dalmatian coast .... 

Palestine coast 

Rest range in Cyprus .... 

Caucasus , 

Southern France at 1,000 feet , 

Syrian mountains 

Generally speaking these conditions are more continental than occur in Aus- 
tralia, hut the coa=tal localities can be more closely paralleled. The Dalmatian 
conditions with respect, to mean and amplitude only are reproduced to the north 
of Cootamundra, Sew South Wales. 

The Palestine coastal conditions are reproduced fairly accurately near Gerald- 
ton. Western Australia, as indicated ; 

Geraldton, W, Aust 

Chapman, W. Aust. 


It may be convenient at this stage to summarise the mean temperature 
characteristics of the habitats of the pines under discussion. This is set out in 
the following table. The summer and winter temperatures are obtained by 
respectively adding- and subtracting the mean and amplitude for each species. 

Table 11 

Summary of Temperature Conditions characteristic of &fcan Habitats of 
Pines of the Mediterranean F.nvironmcut. 


jR. radiata - 

P. canariensU .... 

P. pinaster — Atlantic environment .,,. 

Mediterranean environment 

P. nigra .... - 

P, halcpcnsis 

The main contrast is between (hie maritime climate associated wiiti P. radiala 
and the continental climate associated Avith P. nigra. ¥a: the first three species 
listed it is possible to reproduce fairly accurately the appropriate conditions in 
Australia. For the last two species the amplitudes are not reproduced in 

The temperature homoclimes of the first three species are given in the maps 
of ftgt 3, and fig. 4 } 5 and 6, 

The first introduction of pines was for aesthetic ptirposes and for shelter 
breaks, and it was not till the very end of the last century that plantations began 
to be established to yield softwood for industry. 



n r. 


.Summer Winter 
° F. ° F. 




64 52 




66 46 




60 49 




72 44 




68 32 




77 43 


The fir^t successful introductions were undoubtedly rhe Mediterranean pines. 
One winch, owing- to its economic unimportance, has not been dealt with climati- 
cally, wa> the first favourite aesthetically, the stone pine {Pir,;ts pincn). Its 
umbrella crown and its edible seeds contributed 1o its choice. Next came duster 
pine and then Aleppo pine hi order or favouritism. These three have survived 
in city gardens and park1auds v homesteads in the country and in windbreaks 
around thy paddocks, fjftfli Pmh to the Northern Tablelands of Xcvv South 
Wales, wherever the rainfall was sufneieni and ihe poil suitable. Canary pine was 
a later introduction, and it was not so extensively planted for ornament. The 
seed tv& hard to procure, ami also the transplanting of the seedlings is trouble- 
some, I'ilack pine under the name of Luritrto was. planted, but only rarely. Growth 
figures of some of these species planted for ornament and shelter are of interest. 


It was not 10] the seventies of Ta?t century that this pine was introduced from 
Cahfonnn, Its extraordinarily rapid growth nude it a prime favuitriic. and the 
Mediterranean species came to be neglected. Grown as a specimen tree,, or in 
avenues, it puts on as much a.i three- quarters oi' an inch in diameter and as much 
ns six feet In height in a year, h is very rn-w to rmAc in rhe nursery and can be 
planted out when only a year o 1 d. It was only natural that this piue should have 
become the prime favourite for planting, both for ornament and shelter. It was 
planted t vcrv where in Australia where trees couSd be grown at all. and it has 
survived m the belt of winter rainfall where this is more than 20 inches, it has 
failed in the summer rainfall befe, and its northern coastal limit in New South 
Wales m aromid Taree. In the summer rainfall and subtropical warmth it develops 
a vcrv br:mchy distorted habit of growth ivvA succumbs: to the attack-) of D:f<!on;' 
following injury by summer hai'noniv. Aontwvy pine & eot a lone; -live ! tree. 
i'Vi»}fci-e4 M i rfi B&*K V' 1 '"* 'AA-k pitted for -4Mj y*fcrj«. '•- is very pJiOrt-dfwd, 
for it only attains about I2G years in its native Mte in America, ihe introduction 
of this tree h too n cent to permit A m eAr^-ne of ii's iife hi the &»st cA'kA-* 
environment and eA'pbk siluatAn. fj ; : :;lrer.dy ecvlaki ih;>\ h? life is a very ?V>0Tt 
one indeed where rain is dehcient and -u! .-<uAhA\s are itfjfl tfirrn Lfeicrv. hi these 
circum^anrcs the !tfa? 6f of CT*;fu , CH!:i' -peci , .:r; :.;'. p.£ ! awrue rhr?5 & Ln£f 4 ,vt*la $J an; 
46 years. All round the h?;t:r raAAd are-, of southern Australia may be seen 
graU't specimen:, of Monterey pAe p{ 60 ye^rs o!d_ Some of Ae uoblesi of I Ann 
have h^evi feiled in rcetva ?pdM n\v\n- to the shortage o* softwood caiued by Utv 
war. Specimens four lev! A dfci vter :ivx, I2y r^=t in height are stiii *.o be fu'uvJ 
in :hr- iknvnd (fr$rie£ of hAv; South WfcWi frtefrfj? A>A vain and ftjft eo-- *-.:;.'■ n- 
are vcrv favourable. Sreh tine trees *i*9,v be -ecu in ad the saitfeMW :Aates 
includiiu- Western lustra. 'a, wkc-o the ram fad is more than thirty ruches. As 
the ram fall drops the rat-v of r:rcw:h etv! tie longevity ju; reduced, Whk altkrai* 
c urner-ating in some measure ihe.rc is an improvement, and very big Ivtc^. 
60 year; oiu. are to be found on the .sew South Wsdes tahhdands. 

Aror.: I Adeiame this piae e {s'flj 24 A; |- T ^, fti iha-;e(cr and ^10 f^i in height 
li? 5'M- 1 ) '^0 \ears. In rifAcri. ftettpippj ^roden.; there are two syediu-ns 23 iu^lu- 4 
in" a;:d 71 a:::' 75 tea io hA?ht. L :s a very eommou ornamental tree 
ar-GLtfrd ^A' : h:K:'T;'. -\: ^miLrr-on, hi the Atlcs ra-iav; CVph.U Territory, Iji-re ate 
;; jJcvlr 'h'e spec : m-n:s 2v ijich::; i>i diai^it-t::r r.v, 1 70 years old. It is a rarer tree in 
j£e c-oul' :e:. ( io;,-' r.: ; Xow A -iu.h W:;A^ ; ( -A p'v.-> {]\ fyutfif rift I tio>5 l--'*t L "- 
and he;;; upAe':<I it in mM;y! ,. arA honKv:--;i-:s. 


Cluster Fink 

Around Adelaide tins pine attains a dimeter ttf oQ inches arid a height of 
80 feet in 50 lo 60 years. In tite titji TVfe jn ljtob&t iliere are specimens 
40 Inches it. diameter aad up to S9 feel high. Very i>oad specimens arc lo be seen 
in the better ram fall areas of Victoria. There wAre some magnificent specimen? 
at Duutroon in the Australian Capital Territory, one Was 36 inches hi dbmetei and 
70 feel. hi^h. Iu Use coastal region of New South Wales attacks of Chcrnie* 
reduced Ihe vigour of this tree. 

1'laci; P;ne 

This pine is rare as an ornamental tree in Soudi Aus'.nlin. In ^ ictorja :herc 
arc fine specimens in l.kil'arat, jjfiUht Macedon fold fa many oi the parts ni SlittilfF 
towns. Some particularly flourishing trees are to he sem m the mountain vavtv- 
ships, svtcfa as Yuckandan, soir.h of Alhnry. 

It h tiot common 51 New South Wa'es. though it 1$ to be found in mitty oil 
the stations of the southern highlands. Two {rood specimens are to be -ecu at 
Kupperkumbalone; in the Australian Capital Teintory. At Wa lennvan- ^.ere 
is a tine row of them, and they are so well acclimatised '.hat crops of seeming 
have csrablidied themselves in the adjoining paddock, h ones not appear lo ha\e- 
been imroduced ill Tasmania in the early clays. 

Aleppo Ih^ii 
Aroimd Adelaide this nine attains a diameter of 30 -inches urul a height oi 
70 feet iu 50 to 60 years. It is frequently u^ed m a windbreak, iu many inland 
dry areas, especially where there is lime in the soil, thi- tree flourishes where 
cluster pine E&51& At Murlgec, Xcw South WeIssj it is the best street irce. 

Summing up the position s£| far as the oruameuUd a^J arboricuhuntl planting; 
goes, a 1 the species of pirns rifting iuvesrigcaed have sr=ereded iu all ; ■;-:: ^ of 
southern Australia where rainfall siitliced and %fni eouduxns permitted, k must 
be remembered that trees planted foj* ornament, avenues rod windbreaks, iia.vt 3. 
much better chance of survival than Iras- planted in the doge formation 'nerc^ary 
to assure a crop of timber yielding in^s. The economic plantation of tbe forevter 
affords, therefore, a much better test of the suitability of a species to the various 
factors of environment. 


With her apparent weahh of timbers, it was only m.iural thnt the nrtmci&l 
creation of forests by planting should have come very late in the history oF Aus- 
tralian development. Tl:e first Slate to find the rued, for growing- timber was 
South Am.traba. which wan deficient even in hardwoods from the beginning of 
her iuVm-'v. So in 1888 the first plantation;- of -softwoods were established. \W 
that fla'ic the viijorotTS tfrttwtli of .Monterey pine had been recognised, audi in con- 
sequence it became the- chosen foresters' tree very early. The areas planted each 
year were small, and. in addition to Monterey pine, some of tbe Mediterranean 
pines v. ere also planted.. In some, maces the ffpfefefo£ were mixed and valuable 
data on comparative growths were obtam'-d. Wirrahara, TUmdaleer and Kuhpo, 
also Penela mid Mount Mehrtyrc in tW South-IIasi. wen- all sites of phmtadons. 
The rate of annual planting was gradually increased until the twenties -of this 
century, when it readied several thousand acres a \ear ami consisted a 'most 
entirely of Monterey phie. ily this tiir.e. too, the other States bad embarked cm 
softwood plantations, so that areas came to be planted in widely separated parts 
of the continent. While Monterey pme was accepted as i\ e best tree of ail. uther 
species, including' the four Mediterranean pines, continued iu be tested tinder 
plantation conditions. 


Monti-:>U'v Pine 

This species has proved quite successful in the better soils in the karri country 
i if the suiilli-webt at Western Australia. Areas of suitable ajtil were, however', 
sft restricted that it was found best TO make cluster pine l he main planting species. 

In the South-East of South Australia, plantation-; of this spcc : es have attained 
the h$st dbv^uprricttt in the cuntincul. There are now over 80,0(10 acres 01 planta- 
tion in this region, and the cut of lop's amounts to around 5.000.000 cubic feet nf 
timber. This supplies both the sawdoys and IhiuniiiEjs for the production oi 
cellulose*, also logs for the making of veneers. Some idea, of the rate oi growth 
in this ideal climatic site will be gained by a study of the yield per acre. The 
maximum rate of growth in cubic feet per acre is 500. The average is around 2/5. 
8.000 cubic feet of logs rntve hern commoniy cut per acre from plantations 26 to 
30 vea^s oM. The success of this species m this region has led to the formation 
of private coirrpanies. and these have established plantations adjoining those ol 
the State. Department o[ \\ Uods and KoresU. It may now be confidently accepted 
that this region will become the inosr important softwood forest area of Austral.;:, 
and that this is wholly dip to the. success of Monterey pine. 

While growth was slower, excellent crops fotve been grown a-1 Hundalccr. 
Wirrabara, Mount Crawford and Kuitpo. Compared -with South Australia, 
Victoria lias not planted so exten^ivehv The h'_*d plantations are in the Rrkdli 
ciistriet. The growth here \-. not so good ns in the South-TLa<) of South Anslrallu. 
There are a number oT small plantations of <^reat interest, but nowhere is I: hjere an;, 
large body of pine forest. The 13a!latat Water Supply authority established a 
very s;iecessful plantation in its catchment area. Private companies have made 
plantations, and ihe one ai Portland is the largest, i he growth approaches that 
oi the Mount llnmbier district. 

In New South Wales Monterey pine has not done well in the coastal site-. 
On the other hand it has succeeded in the mountain iev;ion in the Tumut disrrict. 
Here an excellent rate of growth has beat maintained. It is unfortunate that the 
planting policy established in the twenties was abandoned for 14 years. As ;i 
result 'che total area of plantations of pine is Insignificant. While there arc many 
small plantations, nowhere is there a sufficiently large area except in the Tumut 
district, to test thoroughly the suitability of any of the species- 

In the Australian Capital Territory, pine planting was started in 1914 fur 
aesthetic purposes to cover the bare St rondo mountain near die capital. Economic 
plantations, chief! \ r of Monterey pine, have been established since 1926 in the 
higher rainfall areas in the foothills of the mountain range widen forms the 
-western hound an of the Territory, The growth has been satisfactory. It com- 
pares with average sites in similar country in the Tumut district of New South 
Wales, The rate of growth is less: than in the South-East of South Australia 
and wnnld average 200 cubic feet per acre pet year v In spite of the dryness Of 
the Strondo site the aesthetic plantations have done quite well, but although veiy 
beavy ftiffrruixg has been carried out ihry wouM seem to be reaching their physical 
rotation and it Is doubtful whether they can be carried beyond 35 years. This 
area is a good example of the economic value of Monterey pine, Strom Jo has 
supplied the bulk of the flooring for the tfoftaged in Canberra, and, in spite of the 
fact that the climatic f acU rs are definitely unfavourable, no other species of tree 
would have given such returns. 

Small arboreta, established by the Commonwealth Forestry Bureau to test 
the v of different exotic trees for plantation purposes., gave very uMerec,- 
i>£ results. They were established at altitudes varying frum 1.500 to 5,000 feet. 
Up to 4,500 feet Monterey pine proved the mos: vigorous of all trees tested, 
In I he plots at 5.000 feel, it failed to become established w^r three Micecssivc 


seasons. It was not clear -whether it was tlie eiiect oi tong snow cover or of low 

In Tasmania, only very restricted plantations of ibis species have been made, 
and they have noi been successful. 

Caxarv Vi^r. 

In South Australia 2-10 acres have been planted and they are scattered in areas 
at Wirrabara. Mount Crawford. Kuitpo, Second Valley and Bundaleer, Tt has 
clone best of all al KtiUpo. where there is a 4.1-yenr-uld stand. The trees are up 
to 22 inches in din meter and 90 feel in height. The oldest individual trees are at 
Wirrabara. and al 65 years are 28 inches in diameter and 104 feet in height; 

Western Australia only possesses experimenlal areas of this species up to 
8 acres in extcuu These cover a wide area in the south-wcrit id, the same localities mentioned In connection with the Aleppo pine, With the addition 
pt East Kirrup. 

TIk* largest trees arc at Iiamel, nn:l at 45 years are 15'!, inches in diameter 
and So feel in height. They have done well at Manjirmip; there they are 21 rears 
old and have thrown to an average oi EG^j iuchet. and a maximum of 14 inches m 
diameter atid teigltts of from 70 to 80 fad. The species has done well at Naunup 
and Helena. 

In Victoria there arc uo forc*4 plantation* of ibis species. 

New South Wales has used this pine for reclaiming countrv in process erf 
erosion by wind along the Murray at Moira. it is even healthier looking than the 
Aleppo pine with which it is associated and is making vigorous growth, 'Tt is 
exceptional to see a plot thai is not a uniform hcahhy dark g recti colour,' 1 reports 
the Forester. The mean annual increment in diameter is 0<4 inches, and in height 
1-8 feet. The maximum figures arc 0"6o inches and 2'1 feet respectively. Unlike 
Aleppo pine, which stands the {rents of the hollows in this district, Canary pine 
js suscqnible. These difference;* in frost tolerance are in keeping wuh the differ- 
ences in the climatic associations of the :avo species in their native habitats. 

Li the. Capital Territory only a very small area has been planted with this 
species. 1' gives promise at Stromlo of making- mill logs, hut canno;,, of course, 
compete with Monterey pine in vipewr ol growth. 

Cluster Ptne 

In South Australia areas of plantations running' up to 1,500 acres have been 
established in the Kouth-Ras!. The earliest trials were, however, made, at 
Bundah'cr, where there is one acre sixty-one years old. It was also planted at 
Wirrabara, Mount 'Crawford, Kuitpo and Second Valley. The development 
everywhere compares favourably with that of ihc artificially created foresrs in 
the Laudes of Cascony. Whcrr it was planted in alternate rows with Mon.erey 
pine it bas been suppressed by the more rapid grower, cxcepl on sites where soil 
condition* have been unsatisfactory for the latter, It premises to play a larger 
part in the planting programme of the Si ale, as soil surveys reveal areas where it 
is likely to be the. more successful. At Wirrahara il lias attained 93 feet in 65 
years and a diameter of 30 inches. 

Jn Westei-n Australia this species has been accepted as the best pine for the 
coastal p ? ains from Perth to Rnssehon. Considerable areas have been planted, 
mid ai first a good deal of trouble wa.s experienced. The growth ni some areas 
was very poor indeed. Research into the cause revealed that it was a question of 
soil fertility. The use of superphosphate has, it is claimed, corrected the trouble 
fKessell and Stoaic (T>38>, Terry ! 1930),] Considerable work has also been on the question of the different races of cluster pine and Perry (1940) 
reports that the type from Leiria in Portugal wr.s found to be the most vigorous 
and save Hie most svmmctrical share of trunk. 


hi Vieroria the Forestry Commission has established some 2,800 acres of 
plantation at Anglesea. Franks ton and Waare- The oldest is now 21 years, and 
diameters are up to 14 inches and heights 50 to (35 feet. It is claimed that this 
tree has been moderately successful only on the best sites iij these areas. 

Itt the Australian Capital Territory it has only been the subject of experiment 
in Lhe altitudinal arboreta. Tt l)it£ not clone well, but it is interesting to note that 
il has not b'en killed by the. very severe fronts that are experienced in this territory. 
It survives even at 4.500 feet. 

In New Wales it was plana J in various localities between Eden and 
Novvra. Nowhere have large areas been planted, so that it cannot be said to have 
been serioush tested. The interruption of all softwood planting was a contribut- 
ing factor to the lack of evidence in Or- State. In the south constal plantations 
it has not proved successful. C/?r;. ; :/cv iiudouhleudy affected :1s i.ariy ^nrwth. 
No attemp; has been made to select the hem race for the region. 

In Tasmania b&ty small experimental plots have been planted at Strahan on 
ills west coast and at Triabauna and Hasting on the east coast. They are all 
very y-ttfiiig, but reports, do not give much cnconra^emenl:. The Department has: 
now tn'ruhteed the Portuguese nrain, from wide!) better results are expected. 

Ib.ACK Pl.Vh 

In South Australia the total area planted is about 200 acres scaitered fa small 
planrar.ons at YVirrabara, TVlomu Crawford, Kuitpo and Second Valley- and i!h 
the Sotilh-East at Ptnohi, hYlyora ::nd (hanunc. with largest area a I Mount lhm\ 
97 acres. At Wirrabara it Ira- reached 82 feet and lb inehes diameter in 65 yt-ars. 
Xowheie is it at ait comparable w'th tee gr&wth seen in E;u-ope. 

in Western Ausiralia planting has been confined to experimental plot* in the 
ioca'.i'.:e> indicated under Aleppo pine, an; in addition at Pcmherton in the south 
of the r&iYi country, where its growth is reported as good, in ail other 'ocale.ks 
the species has done barij . The trees are AS Let and a diameter at 
5' : inches at 1/ years at Pcmbenor, 

In Victoria some 4.300 acres have been planted in the following localities: 
Ahv Vai'ey. Aue^esea, lM^hr. Cre^wrck. Tdheei Weir, j/iankston, liccchworilt, 
Xarbeibong, Ovens, Str-udey, Taoron^Ei, \Vreau\ It :;as on'y sneceeded hi regions 
\v\u:n f'e nenfall is; over 39 inches and vdiere the soil i* of good class. The oldest 
areas a<*e -12 year.-- old. and the he 
the diameters from 8 to 14 inches. 

Jn /sew South Wale-, this -peele; \\ti* trk-d in :u\ the e: pcrhix ntal phiiure- 
liens, h ■- V. ; -:t growth is at Pey/,0. Na'ho and Loud 1 M:ue Foiv^t iver tie suaih 
border. These, are all e,ood rahuad and Jitgli sires. There is a falling oil 
f,,' g*eevA i:i Ae drier r£::\^^. A fnsl was made with -^eed teem the (. alabrian 
forest H Ira'v, and the yorme. live- pfaptf out at Pdet I Hit *Wftr Hallow In the 
mouulaii^s, They are only 11 years old, so fert it is arjt possible, to say nu;eh 
about their iirowfh. They are heathy and the survival h 99%. The height is 
2\ feet and diameters r;re. no to / ;ur' ,: ine'e^. 

I-i the AuslreAm C apiied Terrier, y an area of this species sees phm;ed [*i 
Fioiecs Creek In rhe fool) if el -t of the raw- 1 he seedrdi Cfcifc: t'rom ! ersica, but vt*vj- 
mixed races resuded. *il:c best type has mj:'e fair tre^ bu: thee arc numerous 
rUmud i-o's vrltii 4eet ueed'e-. In the a>i^ud : na! evpcr'meetal \,]U^ already 
re'err:d lo. r!:r^e races are under ir'r/:, t'~e Lalahrhin, the one from the Cevenues 
HrWftfc es i\ ?**$?& v:<:\ £#hn\%iiW, a: d the hes( Coiv-ni type, ddi are heaid^- 
aej VJgWtfrli. Or the p!ai:is (■( Cauli. -rra there are two pfots ol the Corsiean 
i vpe They have made ireful trees in dO years, hut are no; --.ufficien^v vieorou^, to 
encourage further planting in this drv a'X'a. 


Ai.kpfo J-'ixe 

South Australia again leads hi area planted, but the tola] is very small around 
230 acres :fl all, divider! between Wirrabara, Btmdnleer, Kuitpu and Mount Huvr. 
The best growth was attained at the first two stations. At Ruhpo it is only fair, 
and at Mount tturr it is poor. The maximum height growth was reached at Wura- 
hnra 101 feet with a diameter of 37 inches. 

Iu Western Australia it lias on.y been planted m experimental plots. They 
cover a wide area in the south-west. Mandating, Appiecross, Hamed, Man] "uuip* 
Harvey; then baric from the coast at Donnybrook, and Nannup and in the karri 
country at Manjinmp. The hest resu'ts are recorded si App!eetoss near letth. 
Twelve aere^, now 15 years old, show an average keig'ht of -M feet and a diameter 
of 5£ niches. The trees are vigorous and of g<nd form. A small p'ot of a third 
of an acre at Mrtn daring, now 24 years old, shows a mean heigh: of 52 feet and 
diameter of 9'2 inches. Some of the break trees ruii up to 111 inches til diameter. 
In Victoria experiments were made of planting it at Cvens and Itecch Torest, 
hut i'; tailed in both sues. 

In New South Wales this is one of the jpceie^ chrsen to arrest die wind 
erosion at Narrandera and at Moira on the Murray, Mr. ih U, JAvles repoAs as 
follows t 

"The plantations at Moira show the best growth of any Aleppo pine 1 
have seen in Australia. They are growing' on sand dunes previously timbered 
wirli CoHHrls ghuca. The tftes are healthy and 1h? stem-form remarkably 
g.ew.1 for the species. From mi mory T would say that these plantations are 
ei|ual to. {i nor better than, anything I saw in the south of PranecA 

Its average rate ot growth, is 0*37 inches in diameter a year wul: a maxim t m of 
0*64 irehes; the eorre^oadinp; annua* height increments are l\ feet and 2 feet 

Tun (Aov.-rn oi- _\ioxr:ii'E\ pixii t.x KaLatjmx Te Kxvej >;;?■: i:\t.m, Or-;:,v, mxs 

A sti:Ay in a numh-r n' _#ea.Mt*cK m southern An-; nib of the various aspects 
of the raie of eriweth <>f Monterey pine has been erAe! c-ut hy the Corrmou- 
weahh Fore^y I ri-ci, Tie nxards of svai-.e^ai eAu;evt : on o" shoots gkjnvcd 
that ai MeuiU lmn\ m SrtfcJJs Ar^uahA, gmwih starts- Jij uiid-Au^ust. and flfn 
i- pvu on between then and Arc* mher; inert is v>,:--{rk:---\ growth in \Yil ten rme;\ 
1A;A end fettWtett s^1*fjl h£-> jnne itffi ot the :>i:vx\' A\tm fe akh I Aewrth 
continue--. Ai t- e y^::r carer: tor A : ; \: to {our week-? in July. in hew bk-W-H 
\Yale-\ wke.'r V':' si^tior^ were under oAcvvAiotr: rt -:■ ■ \ A summer ralirfnd ..reas. 
t'M*% A ilk- -rewth wa. per on hi A. sp'Ar: a-A ciltJj r;:i;i:);cr.4^ fc-1 6> A A- 
suavex r v> ixhs an": aronnd SS^g a;'u:!. -A;. : :" M i ere.\'/th uer^ire | ifi fitly- 
In x.zirrr: -tu:vHCL i;^'w0*' stiriors :.;■ i:;Uch a^; .^e-A' viK'i pijt (m iu snmn; - and 
i-if^^ ai ;vm;j:. in t'".e cry -j'ni \*. e.; L'-mbei'ra, 4J>; or the 5 Aei 6 irtehv- rtnuual 
^Vjg^ o.eveie>pr 1 he.w en Seiuuidvr an ' XuveuJ-er. \2 L r feart'S4a O^emii r ;u..l 
J:-nr:;ry. loy^, r : eL\veeu Ma-ch an-:. A.;:r;'. atvl 2^>v A ?,;:o r :i1 d jutic. Xo '. e.wh 
vrs reeor,led >n \v}y ;■!■- /in-nsr. Tla; itr^reineei A q -a oi t/.e trunks ■-»' en 
aere i?f irises ?n *.he S'^ne ktcrihy Aovtd the toih win-: 

Pt-ra - >;<>y _ 9-i ? Tcr' or /V% 

[•re. - JAfi ,. (J-o ., ., u '>% 

^.Ateit - A ; .nl 0-9 ,„ „ .. £% 

JS*i3 - June - - 0-g t! ,. ., 77v 

The fbWit irrovtii e: ^AniAr'e;; in W'er-t^r;: ' :'•: .ehia showe ; u vnvrnAu 
fiY>m other stations. Growth started in July, and 27% c\ the annual sheet was 
put on between then and the end of September. 16% w y as put on between October 


and March, and the greatest growth took place in the autumn with 57% corre- 
sponding with the heaviest raim 20*9 inches at that lime of year. 

Deuclro.;rraphica! records show that dtirtig periods of drought not only 
docs the diameter increment cease but shrinkages occur. The same phenomenon 
wjH observed hy Dr. MacDougal hi the forests near Monterey and Carmel. The 
ciTfcreuce is that In its natural habitat the drought and shrinking occurs in the 
autumn, for. though Title rain falls, ihc misty weather in summer reduces 

transpiration losses, wh. 


Australia shrinking has only been recorded in the 

summer mouths, and there is a significant increment in autumn 

Mdlett of the average increment 

were kept waterer! to oijmmaie the nun I a i 

Monthly measurements hy Mr. Bo- %. O 
in diameter of ten trees at Canberra during the year 1 Q 43 have been plotted in 
relation to temperature and cTmniic conditions in the diagrams of fig. 7 and S. 
Growth is seen to he restricted by low temperatures in mid-winter, and by low 
rainfall in summer. The climatic index used, the ratio of rainfall to the Ovth 
power 61 evaporation | PrescoU (VJ4S)\ f permits ot the assessment of the 
effectiveness of the monthly rainfall as controlled ]- } y the evaporation. A limiting 
value to this index greater than approximately D'8 is indicated for the period of 
effective rainfall. The greater temperature nm#e encountered at Canberra may 
he compared wTh that of the Caliiornian home of the species, the mean summer 
raid \\:::ter temperatures of which are indicated in fig. 7. 

In tig. 9 is illustrated the growth cyele of this pine in relation to mean air 
temperature, based on observations h_V Mr. Millctt at Yarralumla, A.C'I' ., during 
the years 1940-19-44, on trees that 
factor as much as pos- 
sible. The increments 
recorded are the tour- 
weekly averages for 
five trees. To he noted 
are the much greater 
growth in spring than 
all the autumn, and 
the absence of growth 
in the seventh and 
eighth periods in July 
and August. 

The experience in 
'die AtistrahYm Capital 
Territory aur! on the 


+ 2 


outhem tablelands of CJ 
New South Wales 2 
surest s limits of 
:olera:ce of ibis pine 
ovdstdt,' the rather re- 
stricted ran%e of irs 
native habitat. The 
r'mii in die Territory 
a* A 500 feet is asso- 
ciate! probably with a 
im-au annual tempera- 
ture rjf 4? c F. with an 
r.mplifndc fet 12'5"F., 
corresponding to a 
mean temperature of 

1 (g) 

i © 














(S) _ 



1 IS! 



1 a 










40 50 60 70 °F. 


Fi-. 7 

N.K'jn iiicr-cment tfi millimetres per month of ten trees or 
Monterey pine. 2(> year* -old, in plantations at Canl-erra. 
A.C.T.. in die year 1!U3, plotted agawst the corresponding 
monthly te*npc -ntures. The dau arc tlipsfc of MitleU. 


the coldest month of 32° K. and a mean tempo rati ire of the warmest month of 
57° F. Assuming that 42° F. is the mean monthly temperature limiting growth 
ami a probable rainfall of more than thirty inches, growth will probably he 
possible for most of the seven warmer months o," the year with occasional checks 
through summer drought. 

Conditions in the Tumut district are associated witn a mean annual rain- 
fall of 31 inches and a mean annual lemperature of 59" F. with an amplitude of 
13*5" F. Fehruary k a drought month in terms of average values for rainfall 
and evaporation, and November, Deccmher and January are also likely to be 
drought months in a proportion of seasons. A mean temperature of 45° 1\ in the 
coldest month is some 2 to 3 degrees wanner than Canberra, but still 7 degrees 
cooler than the native winter donate of this species. 

The Cuoici: ok Appropriate Spkcics yov. Southern Australia 

The very high yields given by Monterey pine make it the foresters' choice 
wherever k will grow, liven though the mean annual increment drops to 100 cubic 

feet per acre in some 
/t 7 localities, it is doubt- 
ful whether oilier 
snec:c-s would yie'd as 
high returns in such a 
short time as this "re- 
markahle pine'' ,, ' ) on 
ihe same site. 

Foresters fa all the 
southern States of 
Antral in must con 
thru e i o experi ment 
r; wit h i t wcl 1 beyond 
0-0 < tUt regions of optimum 
X china-:;: factors shown 
oil the map of tig-, 4. 
It is a caniar-caily 
tolerant species and 
has shown itself very 
t irmly in the heavy 
frost districts of Xew 
South Wales. Tins 
opens up ;1 very 
large area of moun- 
tain country in botn 
Victoria and New 
South Wales. Rainfall more than temperature iimits t# distribution bc-youri its 
optimum sites, but seasonal distribution of the rain plays an important part. 
Caution must be exercised in trying- to establish it in tl e regions of well distributed 
rainfall, while to try and grow it for economic purposes in the summer ra.ufa.11 
areas i^ to court failure. While 30 indies and over of rain are desirable, such 
excellent results; have been obtained in a wintet ramfall of 22 to 25 inches that 
foresters should continue to experiment with this pine in these comparative!}' low 
rainfall reeions, low, that is, from a forester's point of view. 


10 X 


0-5 — 




Fig. H 
The increment &$■{$ Of &S- ' plotted against the monthb 
climatic ftroftX of moisture, conditions. The indrx is tin 
ratio of precipitation to the 0-7ih power of the evaporation 

<"> One of the botanical synonym* ol ffirs pine is P. iusiynL\\ and in the early days 
'ill its introduction in Sooth Ati$rrath ft was officially rcf~rrt<l to in Anneal Report a- the 
"RL-maikable BinFc" 


pine. Experimental planting should Lk_- resumed on the 
best nice of Portuguese pine from die forests of Lc-iria. 



Of the Mediterranean pines, cluster pine alone appears to offer a promise of 
economic return to the forester. h has succeeded we'd outside tti tjpliimun 
climatic belt in Western Australia ahov-n on the map of rig. 3. (l itf& proved a 
desirable tree in the soils of South Australia where these were found unsatisfac- 
tory for Monterey pine. Lr will play a more and more important part in the 
bouth-East of that State, not only in bringing into production these poorer soils 
hut also in breaking up the very large area? of Monterey pine and so reducing 
the spread of disease,, boih entomological and mycologica'l Further planting of 
this species is indicated near Adelaide. 

New South Wales present- an encouraging field for the establishment of 
plantations of cluster 
south coast, using the 
Further up on the. 
tableland the race 
known as var. hainil- 
iowl might well he 
used. So much better 
i? the form of the 
Leiria race, that even 
where the climatic 
factors point to the 
Use of the Mediter- 
ranean races the Por- |— 
tuguese should be tried 

Canary and Aleppo 
pines will till very 
va 1 liable roles in 
arresting wind erosion 
in many parts of 
the lower rainfall 
regions The eco~ 
nomlr return in tim- 
ber from such planta- 
tions is of secondary 
importance, but will 
provide a useful com- 
modity h\ a very scan- 
i il y (ree-c!ad region. 
In the mouniainous 
districts where water 
erosiou lias taken 
place, these species 
will be better adapted 
than any for planting 

on the very poor eroded steep slopes. The Chapman district of Western Australia 
suggests itself as a possible area lor trials with the Aleppo pine. 

Black pine appears to be climatically exot'e. While the experiments with it 
in the mountains of New South Wales are encouraging, there is no doubt that in 
all cases Monterey pine on the same sues would yield higher returns. It is possible 
itmt with western yellow pine (Finns pondrrosa), black pine may fill a useful 
role in breaking up the large areas of Monterey pint in the same way as cluster 
is doing in South Australia. 









/ @ 



40 SO 60 


Fig. 9 

Illustrating the growth cycle of Monterey nine at Yarra- 
li:mia. Australian Capital Territory, based oil the hieasure- 
riicnt> of \f. R. O, MWett. The increments recorded arc 
*lio?e of the mean girth* of four trees over periods of four 
weefefrj for thirteen successive lunar months of the Years 
19-a)- 19-1-4. The trees werp watered to clfnjiriute the raanali 
factor so far as possible. 



Tltti authors wish to acknowledge their indebtedness to Miss C. M. Eardley 
for the preparation of a bibliographic guide to the species discussed, and to Miss 
R. Dow for assistance in the computations. The data of fig: 7, 8 and 9, relating 
to the increments in growth of P. radida at Canberra, were kindly provided bv 
Mr. M. R. O. MillctL To Mr. T. & Stoate. Mr. A. L, Pinches, Mr. A. V. Caf- 
braith. flfc D. H. Pony, Mr. B. U. Jjyfes and Mr. A. Helms of the several State 
Forestry services, acknowledgments are due for assistance in compiling the resuks 
of introductions of the. various species to Australia. 


Bean, VV. J. 1925 'Trees and Shrubs Hardy m the British isles/' 2, (4th Edn.) 

Brooks, C. E. P. 1932 "Le Clirnat du Sahara et de rArabie" ("Lc Sahara'*). 
Soc. de Geographic Paris 

Byles, B_ U. 1932 Commonwealth Forestry Bureau, Bull. No, 6 

Day, W. R. 1945 "Forestry," 19,4-26 

Day, VV. R-. 1946 "Narure," 157, 827-829 

Herrektsck, A. J. 1901 "The Distribution of Rainfall over the Lands." London 

Kendrew, W. G. 1927 "The Climates of the Continents."' Oxford 

Kkksklu S. L m and Stout. T- N. 1938 "Pine Nutrition," Forests Dept. 
W. AusL, Bull. No. 50 

Masox, II, L. 1934 Carnegie Inst., Washington, Contributions to Palaeonto- 
logy, pt. iv, 124 

Perry, D. H. 1939 Australian Forestry. 4, 12-14 

Perky, D. II 1940 Australian Forestry. 5, 85-87 

Prescott, J. A. 1942 Trans. Roy. Soc. S. Aust., 66, 46-49 

Prescott, J. A. 1946 rt Xa1ure/' 157, 555 

Woods and Forests Dltartmext, South Australia Annua! Reports 


AsciUvRSON, P., and Guaebner. P. 1913 "Synopsis Mitlel-europiiischen 
Flora," 1. Leipzig 

Bailey. L. H. 1923 'The Cultivated Evergreens." London 

Battaxdier, J. A., and Trabut 1902 "Flore dc FAlgerie ct de la Tunisic." 

Bean, VV. J. "Trees and Shrubs flardy in the Britisb Isles," 2, 1925; 3, 1936. 

Boisster, E. 1881 "Flora Orientalis," 5. Geneva 

Uonmkr, G. 1934 "Flore complete de France, Suisse et Belgiijue," 12. Paris 

Byles, P. II. Commonwealth Forestry Bureau, Bulls 2. 5 ; 6, 8 

Chtttknoen, F. J_ 1932 Conifers in Cultivation. Repor) of Conifer Confer- 
ence, 1931. Royal Hort. Soc. London 

Cor-TMAN-RocRRS, C. 1920 "The Conifers and their Characteristics," London 

Coste, II. 1906 'Flore dc la France/" 3. Paris 

Daiximore, W-, and Jackson, A. P. 1931 "A Handbook of Conifcrae." London 


Embrrgkr, L. 1938 "Les Arbres du Maroc." Paris 

Fiori, A., and Paoletti, G. 1896-1898 "Flora analitica dTtalia." Padua 

Gordo;, G. 1880 "The Pinetum." London 

Grossiieim, A. A. 1939 "Flora Kavkas." 1. Baku 

Grlnwalo, A. 1929 Seed Catalogue, Wiener-Ncustadt 

FIaurwitz, B., and Austin J. M. 1944 Climatology, New York 

Hegi, G. 1906 "lllustrierte Flora von Mitteleuropa," 1. Munich 

Kent, A. H. 1900 "Veitch's Manual of the Coniferae." London 

Kew P.ov. Bot. Gardens 1938 " H andlist of Coni ferae, Cycadaceae and 
Giietaecac grown in Royal Rotanic Gardens." London 

Lazaro k. Iiuza 1906 "Compendio de la Flora Espagnola," 1. Madrid 

Lindsay, A. D. Commonwealth Forestry Bureau, Bull. 10 

Mueller, F., von 1881 "Select extra-tropical Plants;' 5 Sydney 

Post, G. 1933 "Flora of Syria, Palestine and Sinai," 2. Beirut 

Reiider, A. 1940 "Manual of Cultivated Trees and Shrubs." New York 

Rouy, G., and Foucaud, J. 1914 "Rore de France," 14. Paris 

Senilis 1866 "Piuaceae." London 

Shaw, G. R. 1914 "The Genus Pinus," Cambridge, Mass. 

Thome 1903 "Flora von Deutchland, Ostcrreich una tier Schwelz." Leipzig 




ByR. L. Crocker and J. G. Wood 


Recent advances in the understanding of the climatological and pedological pattern of Australia 
have been paralleled by an improved knowledge of the post-Tertiary historical sequences. New 
concepts of plant geography and ecology have also been developed. It therefore seems opportune to 
attempt an analysis of some of the main features of the development of the South Australian flora, 
and a definition of the factors which have influenced the species-mosaics as aggregated into the 
present communities. 





By R. L. CrockkkC> and J. & Wood ' ^ 

[Read 10 April 1947] 


I I>;TROi>ecrro,v >• . . . r . I 

IJ G?-0L0CTCAt. Factors t r2 

1. Physiographic influences .- .. .. * ► • • -'« 

2. CTmra'c Influences - -• -* S3 

III History of tttk EAm.Y Atstkauan Flora .. .. ,. -. .. - . C6 

IV The Rixr.NT Aridity -• -- W 

1. Agje of Maximum Aridity 1M 

2. Trends in CTniale .. t> + - 1M 

3. General Effect oi Aridity * 1W3 

4. Contraction of Area, ar.d Survival Foci -- -■- 104 

V Development oe Present CoMMTrcmRS .. 106 

L Migration and Re-colonisation .• 106 

2. M gratory Routes -- 107 

VI Types and Problems op Serenes Distkiuction tt -- fOo 

1. Wide (continuous) Distributions , -. • . ..111 

2. Restricted Distributions — Relic and EnUemT- Species HI 

3. New Species and Taxenem c Problems .. ». .. ■• ..116 

4. Some Revision and Further Considerations . . . . . . . - - . 1-0 

VIT Fxor.oCTCAL Ce.NTr.r-TS \HB the TiEVELOi'MKNT oi- - the South Avstrattam 

VWF.TATTOX -• f?l 

\ 111 Discrssiox axb Conclusions * .. . . li"S 

JX SrMMAkV *- ■ r JBJ 


Recent advances in the understanding of the climatological and pedo' 
pattern of Australia have been paralleled by an improved knowledge of llic post- 
Tertiary historical sequences. New concepts oi plant geography and ecology have 
also been developed. It therefore seems opportune to attempt an analysis of some 
of the main features of. the development of the Soutli Australian flora, and a 
definition or the tactors wlrlch have influenced the species-mosaics as aggregated 
into the present communities. 

The following discussion has been confined chiefly to South Australia, hut 
the implications and principles can be applied much more generally, and indeed 
many of them to the Australian flora as a whole. Where necessary, or desirable, 
no hesitation has been fedt in taking examples from outside, or extending the dis- 
cussion bevond. South Australia. 

(*) YVaile Agricultural Research Institute^ ITtiive-rsity of Adelaid 
ft.) Department ul Botany, University oi Adelaide. 
Trans. Roy. Soe. S. Aust., 71 CI), 25 Jul* !947 


Modern flowering plants arc generally considered to have had their origin in 
the Cretaceous. Since then the paUteontologieal record shows a differentiation to 
more modern types, and by the close of the Pliocene most fossil plants exhibit 
close generic and even specific relationships to present-day groups (Thomas 
1936). It is unnecessary, therefore, to go beyond a consideration of post-Jurassic 
times in phytogcognvphkal studies of the Australian flora - 

The historical influences which have a hearing upon the development of the 
Sou'.h Australian vegetation can be considered as being cither physiographic or 

1. PrrYSioo-RAPTnc 1>:flul:xci:s 
( 1 ,» Cretaceous 

The early Cretaceous times in Australia were periods of considerable marine 
inundation, and there was a clear connection of the seas with the ocean to the 
north, both in the Gulf of Carpentaria region, and to the north-east of Brisbane 
(Bryan and Whitehouse 1926). It has also been suggested that there was a 
possible connection with the south, via the Eucla region and the Great Australian 
Llight. If this were so, east and west Australia would have been completely 
isolated in the early Cretaceous. 

Swamp conditions followed in the late Cretaceous. There are, however, large 
gaps in the record, and it is probable that swamps did not follow directly th? 
marine sediments (Davicl 1932). They were, in any case, probably much more 
restricted than the earlier seas. The approximate extent of the Cretaceous seas 
(after Ward 1926) is shown in fig. L 


Fi R . 1 

Approximate extent of Cretaceous seas in Australia, 


The latest epoch of major folding in Australia is also believed to have taken 
place in the Cretaceous. This niuiSL have increased habitat diversity and bad 
important effects upon the distribution of the Cretaceous flora. However, neither 
marine inundation nor the tectonic movements had much direct influence upon 
the migrations of the flora of southern South Australia. During this period land 
connection with the ea^t was apparently continuous, and with the west was 
probably so. 

{2) Tertiary 

During the long interval between the late Cretaceous and the Miocene, Aus- 
tralia apparently enjoyed great stability and was reduced to a peneplain. This 
peneplain is capped by Miocene marine beds in Western Australia and is therefore 
pre-Miocene, and is probabty pre-OHgocene (David 1932), 

The Eyrian series is generally considered Eocene or lower Oligocene. The 
O'igocene proper in South Australia is represented by freshwater alluvial and 
lacustrine deposits, including lignitic beds, and they have been described by a 
number of workers (Brougkton 1921, Mawson and Chapman 1922, Ilowchin 
1929, Sprigg 1942, and others). ITowchin says of the plant remains, "there can be 
no doubt they represent suh-acrial conditions with low relief of the land and 
sluggish drainage/' The deposits are isolated, and were apparently local. 

No major physical barriers to vegetation migrations between south-east and 
south-west can be envisaged, therefore, in the early Tertiary. In the Miocene, 
however, considerable areas in southern Australia were submerged, and this must 
have destroyed the existing vegetation over a large region. The distribution of 
Miocene-Pliocene seas in southern Australia reached their greatest extent in the 
ancient Murravian Gulf. They are shown below in fig. 2 (after Ward 1926). 

Fig. 2 
Extent *ot late Tertiary seas in Australia, 

In addition to destroying the vegetation (he s(#a themselves must have proved 
3 verv extensive barrier to migration, and effectively isolated floristically south- 
east and south-west Australia. Late in the Miocene (David 1932, Sprigg 1942; 
earth movements begam These, continued into the Pliocene, and Through the 
Pleistocene to the Recent, The result was a recession of the sea by the early 
Pleistocene very nearly to its present position, though parts of the South-Kast of 
South Australia were still inundated. 

In areas not subjected to inundation the peneplain which,£ onTle ^ fe lljc cnr '. v 
Tertiary continued to exist and to become further reduced, but volcanic activity 
in the eastern Slates, which began in the Tertiary, led to modifications in both 
soil type and physiography iii spinfc regions. Tbe volcaaism ha* continued on a 
reduced scale right through the Recent, when restricted activity occurred in the 
South-Kast of South Australia (Fenner 1921, Crocker 1941). 

The late Tertiary earth movements were particularly important, in casTCirn 
Australia, where portions of old peneplains were uplifted' to initiate the present 
cycle of erosion in the main divides ot today (David 1932V South- west Aus- 
tralia was less subject to these movements, and the old peneplain, although uplifted 
in part, is still preserved there to a remarkable degree, 

In South Australia the earth-movements were characterised by block-faulting, 
whieh culminated iu the elevation pi the Mount Lofty-Flinders Range system. 
Over what ate today the Mount Lofty Ranges it is likely that the Miocene marine 
sediments which have subsequently been stripped, were very thin, and that before 
the end of that period or early in the Pliocene, plant colonisation of the elevated 
regions had begun. Discussing the Eden-Moana fault blorlc, 5taiie$; (1942) says 
the faulting was strongly pivotal, and, although continuing to the present dav. 
most adjustment occurred prior to a restricted transgression of the sea m |-h« 
Pliocene. The old peneplain land surface is still preserved in certain physio- 
graphic features of the Mount Lofty Ranges. Py contract, the Flinders lack- 
such conspicuous and general evidence of the older peneplanation, and the greater 
degree, of erosion there suggests that the horst block may have been rained earlier 
( Mawson 1942). This is almost certain from a consideration of other £eo!pgft#1 
tlata (Sprigg 1946). The same period of earth movements which built up thr 
Mount Lofty-Flinders system of ranges was also responsible for the initiation (*t* 
the sunklanci of St. Vincent and Spencer Gulf (Howehin 1920, Kennei 19.30. 
Sprigg l'H6) 


The bloekfaulting initiated in the Ternary has continued, though on a rcducf i 
scale, to the present daw A series of retreats resulted in the emergence of \fcrgi 
areas in the South-Kast of South Australia in the Pleistocene (Woods 186Y). 
Fenner 1930, Ward 1941. Crocker 1941 ) —perhaps as a readjustment Lo IxkiK-d 
and dwindling volcanic activity. 

The Mount Lofty- flinders system was further modified in the Quaternary. 
and ihe Gulfs reached their present extern probably in the iate Pleistocene — 
early Wecent. Although ihe data require reviewing critically, it has been suggesLcd 
that al about the same time, or somewhat earlier, regional warping and 
tilling on a bomnu through I ir-dven HU1 - Peterborough and westward from 
Spencer Gulf had dammed bade some of the older rivers,, producing a great inland 
drainage basin (Ilowchm 191.1). 

That the Gulfs have acted as a harrier to migration, from cast to west and 
vice-versa, is shown by an analysis of fickrophyU communities on adjacent penhi- 
sulas of the region (Wood ] C >3Q). 

Apart from lee Gulfs there appear to have been no physiographic bamrrs 
to migration in South Australia during the Pleistocene-Recent. 

2. Climatic 1n*"%uijnces. 

Past cHuudes anil their influences upon Yegetationtil dynamics can only be 
discussed wry vaguely in regard Lo pre-Ouaternary times-. 

Tfaa Cretaceous, or al least the early Cretaceous, fe considered (o have been 
a pptirKl ot low temperatures over most of Australia (David 1032). Il has been 
tiiggttfrgii il iat "the highlands of South Australia, especially perhaps Hie Gawlcr 
Rimjjes, we. its white with snowriekls/' 

Te-riary climates are a little clearer. Apparently Hie early Tertiary was 
characterised by abundant rainfall and moderate tempt rainres, for in the Lower 
Olltfocenc such nics-ie elements ;is \U>lhf>jagus, 1'Hmtfrsia and T*'ista>ta! were wide- 
spread in what are today the rmuc arid part-: ot South Australia (Chapman y§37\ 
Associated with thorn were some of the early Eucalypts. including & Dirnlenu 
Ktrinrr., and E. Klf.^nu, Deane. 

The Miocene marine deposit hehi<r characterised by coralline limestones 
indicate wmn seas, indeed, warmer conditions, than present are considered to 
have prevailed right through to the preglacial Pleistocene ( Whitchouse 1SM0). It 
fe not unlikeiv that, at least during pari ul the. Miocene, rainfall was lower than 
at present. The Pliocene is generally thought to have heen humid, though White- 
house has suggested that in Queensland there may have been two periods ot 
aridily, allernatiug with wetter times. 

It is apparent that insufficient data have been accumulated so far to permit 
anything more, than very general analysis oi pre-Pletssocene climaxes, (hi the 
other hand, post-Tertiary china: cs can he evaluated patter; especially for the late 
Pleistocene Recent, but the evidence s still very scanty and the conclusions tenta- 
tive. The information available has been recently summarised by Browne (]?M5). 

Perhaps tbe best evidence, for a pluvial Pleistocene cbmate is airorded by Hie 
more 01 less extinct riven; and ialces, and the Former cxi-t?ncc of Ia*ngB herbivores, 
in what are now the most aiid parts of Australia (Tate 1879, 1881). 

[) h;is also been considered that the whn-e of the. Picishvene war. a we! y -riod 
in Victoria. (Hills 1938). These opinions now seem to be generally accepted. 
David i 1932) points out that ihe reduced tempera Pares would lower evaporation, 
and this alone would have the effect of increased rainfall. Wfutehouse (1910), 
on the other hand, believed that, although most of the Pleistocene was pluvial 
the very late Pleistocene was characterised by arid conditions in Queensland. The 
evidence, however, now suggests that desiccation was later than ibis. It is inter- 
esting that the Pleistocene glacial conditions, which were associated with semi- 
arklitv in much of the northern hemisphere, should have hapn so generally humid 
in Australia. The reason appears to lie. m the minor extension of the Antarctic 
ice-cap and its stability, coupled with the absence of any major ice-caps m 

Al though rainfall was probably plentiful throughout, temperatures varied 
with Pleistocene glacial and inlerglacial conditions. These changes, must have had 
a mcirloxl effect upon the local flora. Pleistocene glacial condition.- iu Australia 
were restricted to small ice fields in easrerii Australia and Tasmania. These were 
too efttilU f'SW Prooks, 1926) to maintain anticyclones of sufficient intensity to 
seriously deflect cyclonic depressions. The Antarctic ice-caps are considered to 
have been more stable than those in the northern hemisphere (Zcuner 1945). and 
to have had only minor expansions in the Pleistocene. However, whal expansions 
there were probably resulted in a more northerly mean path for Antarctic "lows," 
which, in the absence of any deflection from the Austra jjfti continent, brought 
pluvial conditions to Australia generally, i: would appe-u* emiie logical that these 
continued with only small flnrtualions tlnrou^honl Ihe whole of die Pleistocene, 
inchioirg the huertrbiebil period*;, This luis some sojt.joi I in Ihe geological 

There can be Hule doubt, then, tlat the South Australian Pleistocene Jtotft 
was subject to varying climatic pressures, and migrations occurred with the chang- 
ing conditions. Some time after the close of the Pleistocene the rainfall ska r piy 
declined an*;! a period of desiccation commenced. Before proceeding- to a detailed 
dEcuzEon of the results O; tins aridity, it E interesting to consider the influence 
of the mafor post-Jurassic historical changes on the primary Australian flora. 


The orgm of the. Australian flora has been considered by only a £?;X3iI 
nuu-b.T of writers. Of these Hooker's (U 7 60 i 'Mmroductory FEsay to the Flora 
of Tasmania"'' is regarded a<< a classic. Since then a restricted literature has 
developed, espccEJIy in relation to the genus fwr&Jxptit&t vvbiqji ha.^ been con- 
sidered, as providing rhe key to the autochthonous element. Works of especial 
importance are those or Cambage (1913) and Herbert (1 ( »28). 

Hooker pointed out lhar although the AusrralEn flora was characterise' 1 by 
a large number of restrEred (thnf is confined to Australia) specie/ and 'eiwra. 
it exldhkcd the same primary features as the horns of other continents. He 
recognised four element? as having played the major FG4fcS (ii its development. 
name : v, Ari? r rciie. New XeaEnd-PnlyneEan, ludo-Ahvavao and Australian 
eleiv.ent:-. There can be reduced to three elements-, Indo-MelauesEn, A.durclE 
and AusiraEm. 

'ike Antarctic cEmcnt is widespread in southern Australia and Tasmania 
and erdubhr. cU>££ affiuEEs vEth the lE^ras r:f New Zealand, ."onthcrn South 
America, the rah-Antaretic isles, and to a less eraent Sotnb Africa. The oEeTu- 
tion of this element IhrQUt.Etu: the sotohcrn hemisphere has been he'd as evi- 
dence of irrnn- fanfl c^iuif^on. bo 4 \ht argnnrmts have n$i been '.ouvinEng. 
AtEntEn has been drawn by Gibbs (¥311 ) !o the c'ose aihnitics between the 
Antar itc ekment in Tasmania, and thy Horn of cuaam mountains hi New Guinea. 
Ciibbs considers the AuErE-Artareuc element to have migrated nrlffiliattv from 
fke lai'ter region, and produces menofokiglra! evidence for possihikLes of wind 
dEgersE of t 1 v dE^porcs- Whatever its origin in Ar.E alia, the AutrnvrE c. >::\-i:: 
E ron~EErcd an ancient one elsewhere. 

r Ev b' ;o-MeEnrEr:o ekm-Mu is the Aimiuani one hi senE-'ropEal and Etftpfcal 
\i\ [ ,i A, aiHl A inpor:e_nt in t\\f ilorn tit ihe a:do and semi-arid rcgEw^. 

ChEf EUroet has b._-en [Ecn^ed on the AnstwdEn element which reaches its 
r.iasdm.uii e-vE^pmuu In southern Anvtraka. partionlarly in somh-west AusiralE- 
Jiook-r { . 1 f rO ) thw' pc»:n:r-' nfSI the ricEv s:> Hi ihe tlora of smith-west AusivahE 
in the ckaiaeierEFc Au-.rndhn orders. :grp"uErnnEE sE awenth-, oi vAEh crVn 
!,h E mavmrm in that regEn. the remaEEg one-seventh reaching pt;.-j;,'aio 
dcveEccmen! in somh-eaot EusLrai'a, with none soowdng great dcvido.a:: ' E 
t'.c t:-M,;>. Gardner (lE'2t ha^ indh-ate 1 the diivicuEEs of defining with exa:<i~ 
tuJe \\' 1 A;i--'rEEn eleai *nt. ::i d EArwe r , vh:u muis gMia r :s prL\'EnEy i,..o/E j&yv " 
h; i ■■- *-\Eg to it have loo nu.iu anhiilies whl: tie AnUuvtE ur [vaheeotropjc eErne-]! -. 
e:-d hvh-ed he <n\^-^--'-\-~ v;h ; _d"er tjUW *■- a trr- E.u.o-a'i.a eEntca!. : e d;E a* 
it tav . certAii Tanwho^ r.mJ g.-onv;s \EiEh are eiuire?y Ansirahan. or are ver\ 
poredy r'prc-.uted ehe^here. arc wordiy of ;epa-rde rons'de^a: : r»t], a:rl do" 
rLmnrEd/E deyEo : .'me::r hi yovuh-v, ■-•t Aus'ra'ia nniA have erjiectal signiiicance, 

Ti^e ceiUre n\ origh; of ihe AustraEm eEmtnt E unknown. 1d;e sam-* 
eerera' botamva! fef;>.u:es ar - e Eaw:n in the Anslsahae jva .t--M*EogE;d ocord a- 
rl-ev.Eere. fr-r c;;ampE, the i«r-t dieor> 'h E'>:a nv\ s-^'Ees prjr vv-.-'r . i i;o;: ' • 
E^etacons of ■ larr! (WatEun E>J E gtf-j jv- the Lower iE ; g):or; a i-]'^:'^ 
Ao-irakan :a;ies. witi; sueh genera v.h U:ica!\pfu-: ar.d Bmiks^' wv.s cdEan in 
S-'Uih. An.^ralia ( Eha^ man 19d7). and is ptustttt'dj] also hi the Jigrdte of ^TorwelE 


Yallourn, Victoria (David 1932). 1 fouled- was led to suggest fruru the ordinal 
and gex.evie peculiarity of the flora in ^omh-wesL Australhi tto} \\ estem: Australia 
was the ecmruui of the Australian flora. He point* ovu, however, drat from 
the viewpoint of habitue diversity, one would normally expect h-ast Australia to 
lu be richer in peculiar forms. Bdlo&J <>t the work concerned with the origin and 
development of the AiiFiralinu dement has been concerned with floristic elements, 
and very little attention has been given to the importance, of parallel gcnAgical 
factors in its evolution. 

In any case, such considerations, would have, been oi hide value until con- 
clusions from studies ou the basic geology of Au-araHa bad reached somednm;. like 
stability. Tbc first plant geographer to consider geological factors very fully was 
Diels (1MJM, who postulated the 2 j a>ssibiJi;y of a pan -Australian flora prior to 
its heinft split up by ihc Cretaceous seas. Woo-! ( l p 3(b rJso considered dint the 
isolation of the two centres in south-west and ?ourb-cast Australia., from uhieh 
ndirrateHi has subsequently proceeded, was initialed with the epieomi rental 
Cretaceous seas. Subsequently, it was believed, a !ar^e measure of isolation lias 
been maintained in vvtsi Australia, while considerable admixture with _ ji Jo- 
Mela uesian and Antarctic elements lues occurred ill tbc east. Hooker had suggested 
thai Lite flora of South Australia, which is poor i.i endenres, wuidd be found to be 
tlprTVUtfvje-, and intermediate in character between that of the east and west. 
Wood (Joe. rlf.) was able to confirm tKlS in an analysis of die sclerophvil coni- 
muuides on Krin^aroo Idaud and Ac adjacent peninsula, , and wa» able lu demon- 
strate CMit the South Australian gulfs had acted as a barrier to recent migration. 

However, since Ke>oheiA day tin cniudnchiii atgaments have been advanced 
to supp')n Ins -lecH'-'-diou tlmr Western Australia was the ceutruui of :i;e Aus- 
tralian dement, although it is established as a centre of d^pexbal. In his analysis 
of die dHstributiun of the j*GBU5 Eitcit?yMn$ t Herbert (1928) has conclude 1 that it 
is tjol possible to locate the centre of origin of thai gams, bin. ilia} fl h i airly 
ce'-:^in that it successfully established itself in regions wilb a temperate- clHflalR 
and abundant rainfad m the late Cretaceous or car'y Tvrliary. This paper of 
Herbeiv is also norewoeby in bring the- rirst in which the eilVct of an ember 
aridity was considered sy>U;uiatieally-. 

ir is worth reviewing the eifeei of the geeHXpcal factors upon the dhribtt- 
tion of t!u: primary elements of the Atuaraban flora. 

hirstlv, the significant florf-dic and paleobnUnica! evidence indicates that the 
typical Atislrrd'nn tdem^id probably originated in the 1 Late Cretaceous fit* early 
Tertiary. There is no evidence of its being widespread until the early Teniary. 
r.y thai 2:me. e.g.. curV Oli^ocene, such rrenera .v T'ank^hi, llahca, A\">YwnX 
LoiUMlt-j and EaaJy^-ias were we!I established in South AitsU'nHa* though much 
admixed wdh a W.v ;<eahmd Antarctic element typified by tin: myrtle beech. 
AV-'/Av-z/V'/f/s". Ic is obvious ihat loo ni'.teh atkutton J^g been focused on the 
importance of the early Cretaceous seas as isolating ftrittfiM:^ elements of rhe Vie>- 
ttwifiS-n Mora. In the firs' place, although It is possible \hr,\ vtv-\ and west A^strruta 
were then completely tNulated, ihe : veolo^ical evidence lor I is so scaut that such 
a snppositiLMi must he considered douhCvd. in any case, i^ no evidence n$ 
yet thai ihc primary eiemeni* u f ihe ,'uislralian 'fiora hi.d arisen bv tltc early 
C-retaceote^, and indeed all the avahahle evidence iu- r e;esi : a later origin. H is 
jlrn&abk that th$ seiuhem Au'jirah ; an flora wa-; f$\\s$ in na^mte I'nnn ens'; to westj 
if not through ail the CdTlacetnis dien certainly through die j^rcatrr part of it and 
through ad of the lain-r |ioiiiini. In Smith Anftfr&litf, laud cojmection wiili Semtb- 
P-risL Au c tridia v/as a.ppnreutA coutimiuns throughout. 

The over-emphasis, upon die imporomcc of the Cretaceous epicontintntal seas 
bus been associated with complete nesd-ect, with d;c exeeptieva of a comioeratiou 


pf the importance oT the South Australian gulfs (Wood 1930), of the mairpgr in 

which isolation has heen maintained between cast and west in the long interval 
since the lower Cretaceous. 

Because of tlie incompleteness of the Cretaceous record and the- history &I 
tlie eaHy Tertiary the centre of origin nT the Australian clement will prohahiv 
always trt unknown. \or is it reasonable io surges! tliat there was &n% centre ot 
origin fn*- the whole element. The richness oi the flora of soulh-west Australia 
cau he understood heller in Lenrre of :;u early uide'y-uistribuied flora. Tha 1 a 
large and typical AttBlftlfeMl ehmerq was widespread in the early Tertiary has 
already been established. K portion's of such an element were irub-equendy 
isolate,; in the south-wes* irom general plant gC#j£&i/pfrtetil principles It is logical 
U> as^e-me that with agi? and eoniiiiued isolation a rich endemic tlora would have 
developed. The questions requiring answer are first!}, iu weal period w:^ such a 
pan-AiMrab"an flora likely in have evicted; secondly, Avhen and how did ku'aiiou 
occur: and thirdly, ill whal manner has i-adaiinu been so successfully maintain^ I. 
T.l is obvious that alt three answers are required to satisfactorily explain the present 
distributions, and it will fee necessary ;o refer hack to the potil-jurastic geoioLjicfJ 
factors rmnmarised previously. 

A-: Gardner (1942) points out. the present d^tribmion ot a large number 
of Au^iaiian genera lends definite support to the theory of the jveviou- 
exis.ence of a pan-Australian element. It was ibis fact which led Did.- tfl 
Stigg?-?.! division by Cretaceous epicontinental seas, Reasons have already been 
pdven why this suggestion is untcnab'e. It has heeti shown, however, thaL typical 
Australian genera such as Hakra^ Bi>iiksiu. Fumtlyphtx, etc., were widespread in 
the lower Tertiary oyer a £reat pari of sou:h nod east Australia. I ufortunarr'y 
palcob^tauical evidence does not extend to V\ estcru Australia. X eve rib-; h ,. ihe 
dfcunvtantial evidence is almost overwhelming. It has been mentioned t.!:.u ill 
the fong interval between xhe retreat of tlip lower Co:taecou> seas- and. :he inui- 
Tcnian Australia enjoyed great stability, and during this L i me was reduced t-j 
an almost perfect peneplain, with very few. if any. conspicuous ranges. As a 
consequence, the soil and cbmaiic patieru must have been extraordinarily regular 
and the zones broad. These circumstances must have been very Favourable, ;f 
not: unique, for the wide distribution of the floral units, 1l was ac some stage or 
stages °f ^ s peuepkuratiou that lateririsation, preserved today as; a fossil s •;! 
character, was so general. It is most hlcely, therefore, that the elements wide 
spread in s&tftfl an4 east Australia during ibe Oli^oeenc were also prominent >n 
south-west Australia, more especially :»c these were dines of abundant rainfall. 
FaJltiwrig iateritisalion, and probabh well into the early Pleistocene, soil condi 
tions over much of southern Australia ],n*e*,erved u remarkable uniformity and 
morphological relationship. The evideuee ihen is strongly in favour ql a par 
Australian flora from the early Tertiary. 

This unifonuity would Ottty be presented if no marked barriers to migration 
-were to develop. Should a subdivision of the flora occur, divergences couid suh- 
sequendy more easily lalce place. It so happens that the- lolur period of posi- 
Crelaeeons siabilih was biohen fi] the Miocene by the initiation of epeiroyenic 
earth movements, and by the inundation o[ a considerable portion of southern 
Australia. Although the climatic zonal ion was broad, ana edaphic condition* 
except ionaJU" uniform, there can be little doubt, in view of these seas, that the 
southern pan-Australian flora was restricted to two major regions in the iale 
Tertiary. The erYeet of marine inundation as a barrier to migration w;b heig'n 
ened by the v/armer conditions that prevailed during- the late "fertiarw for this 
ciimatie effect wo;dd liave. resulted in a migration restricting many m T ouq^s to tltE 
most .seaitherly portions. The Mioeencr seas Imd withdrawn almost to that pxtel ■ 
at present by the clo^e o[ the I'iiocene. 


Whilst it is believed frhflt $\i [orbing establishes a s:ron£ case fur i^niittion 

of the Australian dement in two widely separated port ons ot the content, 
it bi Still necessary to explain lipw the isolation lias M&tt more or less maintained 
since the Pliocene. L is necessary to consider in more detail the types of 
barriers which njighl te#c preserved itoJatiuri, They are ot" many kinds. It bas 
has hceii suggested that iiimv or Ufr$ s.imiutancously with the development oi the 
I'erdary kc^s there occurred a somhward migration of the flora in :>ou;hern Aus- 
tralia as a response to changes in ehm.^ic, and I hat ihfe increased the effect of 
the seas Jfl i.-citntlu^ the floras of the tw>> regi.-ns. This climatic, change was 
probably responsible fjlif tin complete disappearance of s^pi? of the more nit? sic 
elements- in Western Australia, like Nolhofa ,'tts, and •which accompanied (he early 
Au^ra'ian flora in the early Terliaty of rioVuem South Australia, It is 
probable that these move m::s : c elcm-ms, or fume ot them, gained suilable niches 
in south-east Australia during th;s southerly migration, because carth-movemento 
(Kosciusko epoch) which were initiated in the Miocene, or even earlier, were 
increasing habitat diversity. 

As mentioned earlier, these carth-moveirieiT s mi Lulled ihe break-up of the 
old penep'mus, and had their mo-::: proiouud cileet m eastern Australia. In 
Wcsixtu Australia Ihcr uj'ihieiVT was slight, and much uf the old pene 
plain is preserved. Ju South Australia the climate was greatly modified 
with the uplift ot the Mount T.ofty~ Flinders Mange, region. Apart from this 
purely local effect the climatic pattern, a>* disrinri from intensity, in South and 
Western Australia has been litrlc cha i£ed. This being so. the arid region at the 
head of the. Great Australian Liiein then, as today, might Ae a barrier to migra- 
tion (A r ui the ^outh-westem province of \\\-t. u rn Ams.raha. With higher 
rninfah* couch' linns, a more northerly ail*l north-easter y expansion of tiic 
flora of soutlt-west Australia would fig logically expected. Such conditions seem 
to have occurred over Ausir.iWa gcneralU in the rieiotoeetic, so that during that 
time llv j \?Aj&?t;i mav r.oi have: been such a climatic bar to migration. 
There are. however, barriers to migration other than those cf climatic and physio- 
graphy. The most in r/ or Lam are soil harricr-i. There is good reason to 
hclicvc that such existed, and an obvious one was the edaphir barrier of the soil 
type developing ^ n the large deposits o;' Tertiary limestone inland from the Head 
rI the Bight. The isolated flora of the souih-wctl was one selected for latennc 
soils and would be largely, if not entirely, unsui ed to sue h different conditions. 
Further, die gflft region of South Australia must have been a harrier to cast-west 
migratioas, as Wood ( l r >o0) has shown. The influence ot the gulfs, however, 
has been too recent to have been of very great significance insofar as the main- 
tenance of die historical isolation of the flora Of the sonLh west is concerned. 
ti is important, but outweighed by the other factum already mentioned. With the 
Oecfi::e in rainfall n?sociated with the early-mid Keceot aridity, there won hi have 
been a general contraction of the Australian flora and further isolation, The 
aridity of the Lhght region has undoubtedly been sufficient to preserve isolation 

It appears that the richness in endemics of the fWa oi south-west Australia, 
especinlh- those of the Australian element, is cspiaincd reasonably well by the 
basic historical facts ami is evidence for an eerly parr-Austra Han flora. The 
pritnarv division between the floras of s;»nth-wesL ami south-east Australia took 
place proliabA in Live AJiocene. Since that time the south-west has been cou- 
linuously, or almo< c<:'miuuous!y, rloiist^cally isolute<t from the south-east, w'uh 
ti.e result that it has pre-tir\ed great florisuA- siahiiily. Jt ]< unnecessary and 
undcsirahle ;o consider tins regiwn as ilie ccm;re of origin of the Australian 
e'.euii nl. bid n is loo-'cal t.) bejicve i;ia: il lias been a c^iiit? of disijersal at various 



Of the other elements it is likely that the ludodMclancsian II a very old oiic 
in the Australia flora, and that invasion? through northern Australia have been 
frequent and sometimes prolonged, It is likely that a major expansion and 
colonisation occurred during the climatic changes in Ihe Miocene, when a large 
scale southward migration is postulates! as a n^ponse to wanner conditions. 
Then: must, however, have been many subsequent rhicfuations, especially during 
die colder Pleistocene. That large portion of the Antarctic element which is 
associated with the mountain plateaus of eastern Australia and Tasmania must 
be relatively recent, for it is hard to imagine its members muling- suitable habdu;s 
in the pem-plained Aus:ral;a of the early Tertiary. l:i all likelihood suitable 
habitat lor tb's SQ'&lVp were no: available until thv onset of the facial conditions 
of the Pleistocene, tor although the he'^ins of ihe higher plateaux may have 
approached those of today, the Pliocene and ore-'d'^ad Pieisaoceue arc considered 
warmer than the present. Gibbs J (hie, cit.) surest ion that thetr successful estab- 
lishment Avar> the result of long distance dispersal must bo consider:! a possibility. 

'the epeieroyeiiic uplifts which raised the main divides of eastern Australia in 
iner casing habitat diveiadtya both cdaphically and climatically, also gave an 
opporiumtv fojr successful canonisations by other canncr.ts^ Similar opportunities 
would be presented by any climatic tiuctna'dun^ of ihe gkuda! ruy.l iiner-gyudai 
Pleistocene. Pa-own (1^45) has suggested a major expansion of the rain-forest 
Payed dtvhi.2,' the Mhnle! - Kiss mid Pleiscoeene inter -glacial. Tin- scenic po,-- 
siblea Although elauatic-induced abeaandou--. occurred in t'-e Xd-bo'Ceue, it is 
neverthe*e?o to he remembered that the ice-sheeas in Australia were so restricted 
as ro b; aauost negii^ible. and the caniate may have been mtieh more stable than 
p re v iou ^1 y tho tight . 

The forecoiir { , r comments on die origin and certain features of the Australian 
Jdora are an mlroduet 'on to discussion on the effects of the pod- Pleistocene period 
pt ar.di." . This desiccation ha--, bad such an important inihiene.e upon ihe and distribution of the present vegetation units and communities that 
it is necessary to consider u in ftrfed detail, 


1, The Ace or Max-Thum Akioitv 

The e:uici nare bf the devo'opniaU of maainmm aridity is uncertain.. It was 
considered by \\ hitch.ouse (1^4U) that the decane in raiutali responsible (or die 
Ctmc systems in south- 1 re-ae:m Oueendam 1 ivourred in t.feb very late JPlcistuccraa 
This vkv; I:jar3 reee^dy beau accepted by Pnar; and Jones (1945) m their strali- 
iTraphaeel oothsie of tin yeoh-adad "ayry of Qutenidrri.l. On die UiUzx haml 
llilh: (bVi-d. from frd important physio^raph:? ^uahts iv. p^ab-wes; Victoria, 
considered thaa The rainfall decreased in the l^ply Keeart, and that tie ivTA o{ 
the ibebtocoiu yca$ relate/ely wvL in Sirajli Australia, neaoio:yeai, :' ( red>'deal 
and \tsi?Wu w.&v&t'-z) da .a have been p,a<etda: 1 \v!d'di ; aar:e-t. a \vtv recaid ::; : y fo' ! 
rhe crteets of maximum apody. h has ivui -^ncye^ted thai the arid prr-a:,. oa.anr<-d 
v 'proia.biy nvocb le: s dum 9.000 year.; aya" < L'tocher 1-;oj, -r.d aaJ. m \,Lxi SoaUa- 
East of South Auslraba the dunedablaim; ci"h:ax arsneia'ed w-Xli the asady. may 
have been a." recetd as /,Gc0 yc:r> ^yo {v'r=ickar an] t uy^a I9-ada 

Tlier- ca!i bj little dauht that th;; :i^yor :.ame •ysrear 1 -, at Aus-rada \C] iw'oilg 
to the one ] criod of chai?-lic -tn.-~ (L'ro:her 1941. Jarowne Iddy. larovne 
(Jar. of,) i» attempiiii^ a post-Tcrdary cin-ouolo^y for Austra: : a has pha;e:i the 
aiiddv at abont 5.0, '0 y ; ars ayo. ddila.a-r'i faiaher ea'dence is -a-'drca. a vaoid- 
seian :hat a j£^cl7ii V^t caa be accejiled, a;;a ft.lvwJiVj.-4y it m: cv !a ha con^iue ' 1 
lu 5 ave bacu about 4.000 b'JMi year^ $g$ t 


2. Tkkxos in Climate 

The most important resuh of ihe desiccation was undoubtedly i&e wholesale 
destruction of die native flora, which led ultimately to accelerated wind and water 
erosion, ]t is obvious that the degree of aridity,, or rather the nature and extent 
of the streets imposed upon the vegetation were entirely rclaih-c to the climatic 
conditions prevailing immediatelv prior to die decrease m rainfall rather than the 
level 01 rainfall itself, for the vegvUndon communities \^akl he in a siale of 
balance ( c y naiiric ) with the edapho-eHmahe envirovmr nt. 1: has heen generally 
agreed that the Pleistocene which preceded the ardhy was a wet period ( sec 
carder), in South Austraha the large tb.iviad'e an .1 terrestrial dtposiis whidt 
cover rO much ot the State are considered to be of tins acre The trend of aridity 
in the Simpson Desert region ha* been discussed hy Madigan (1946). who says: 
"No va''M re:-nn presents rt^c 1 f to justify the widely held view that acidity was 
neater in the [Svjft when the sandridger; had their bud;." He pohtb* out thai the 
faeL thai thtf" >efn T ridi;e-s are now lived is no proof, that it would take a long time 
io replace ihe flora of the pluvial Plcistoecrie, aril that ''tkv ridges eor.ld form 
in the p.sdbly barren t : n:es between the decline of the old vegetation and the 
est.ahhshmevU of die new." 

The relative nature of the aridity is, of course, apparent, but as doubts have 
been expressed to the v ; ew that absolute desiccation ha.; heen greater than at 
present, and became the onestirn is o'" s.ome importance in the consideration of dynamics, the evidence «.i?l be briefly reviewed. So far a? the 
atilhr.V:- can de.erutmc Hicre have heen no suggestion^ that the present-dav climate 
i< mr-re arid ihau thai which led to tie bedding up en die large •YuMratian dime 
KyStfln^r. The question, then, is whether the presem-dav climate is more hiurdd 
than that during the piling up of the -and dune-, and rhinesheeu, or whether mere 
humid eomho'nns have intervened between and now. 

Hil^ (iPd^) considers that in the region of the ancient Murnty Gulf there 
have been relative 1 y we* tee mid drier periods following rhe initiation of the po^t- 
Plei^toceir- aridity- '"'the present ail*! immediate ra~t being relatively- we{. ,? f-fta 
evidence for ivicrmi l Jeni aridity during ihe hend-np of die dimes was based on 
the occurrence of horizon- of s.nneruw-o^cd lime acmumlnh-on within the sand- 
ridge:-;. Tin phenomenon 'm'dn h;t\e I ■'■. e lJii£ to other caries, as Mi^gested hy 
(drneL-r fJWil, #* \ must row he ceresaiered douhfnd- The evidence tor wetter 
cum'dmns in ties reg ; ou. however, is important and is n\ three types. Ph.*! there 
are Ihe "cepi" hills ef the ma dee. T'hese are believed due to ihe sohnion tmd 
re-deposition Ffi duvr-~ of * f -;eed' eryontm with the '>vcV,rtiVMit of powderv cop! 
at the -srtrfro. ami witld tenure a higher rrdntedl thrn that for the ionn.Timi of 
the orhrnrd th'^^--, T!:e inline' tin tit nyt'-nm dnne*^ is a!^)^renllv nei rCnMT^fijj* 
m the rieto;: ^d;^. Seeom.h Ihere rr*- 4 th.e drfffl'nigi" F t ^t tlvouejh the sendddees 
lielow ^.:ih' -\]ha.*tn;ya which are rfriitri T cfti3tl to h-e.e d^ve 1 one,l I^Erirg th-: 
^n-pi 1 ^ b ■•' r* \a riv-^c ih^'ii, e^d ih '-_' t^e a-re't t\£ tee c;e ic-al send divfl- In 
Oucensland, Whitehonse (l^'O^ |»as: made a picn^erre: study of t ?r '--'t-^iiorene 
che-ate:. Vd:d- h:',> *>rcvi ,f ed e ;r'u:n':s m* na a e; r' : nr:oc.h^iea! ale' pah-'-edo 
logical siedn-s e^-^where in An^lnd^p. V<> rirhrh-tct^ •'hea v;h <-h hvbrr.e (Usi sul-- 
<("iue't in jltf utf A re ;^dd 'tt;o;! vdde i fiTrjwe: rl e l^-^^oeeue n!nvial ^iinit. 
d-ere h;i= V-^n a ;- : eht irn-v:-^ \'A i-e ; ef:rh V ;• ; Tnei^al evi It nee '-'.•} hi 
( 1 ) (hgrarf- 1 el-ae-*-- ^m stiSff, i?) th: 1 're^nce >;i ;-_y- <r-.v* ;, ed I'm - ;;.^ ht -n^e.- 
renote }x$ts\ fhe arid %V f fmny ; n vdd'h ihey are ^nj;f|^ in soi's today, and 
(?-\ the redi-!CL ; on o ; da MlVrJhe nrei id ''active type'" rfatffla*. in Sr-nih Ans(r;:ha 
wenr-r cendhir::^ s{tien d;.- arid per : o . were c^r-e'e' j ! (("--.'d-cer l r '41. i'V^) !« 
f*e snge;nsted h.y O") t 4 v wind-!"!- 1 sl!r>-r>-!^. sand- o" d;e S nnh-Ka^t. whi-^ are 
now sunpordrg a ihy sc ! M'o.-: , yM Epregt, (Z) th? ys^ttt^rtte; o! 


A r o!npnla 11 (wivka'n it the lower 7\ fur ray in the lower Ivlurrav which had 
developed a keel as a result of a sudden increase. 111 temperature and/or s^lhtitv 
t Cotton 1935), and (.V) the evidence for warmer sea?> in very recent times ill 
Sotuh Ausiralia, and ?hc ingestion:, that these may have been coincident with 
the. period vi greater aridity (Crorkvr 3946 9 Addi:ioual evidence is to he had 
in the occurrence oi ex*.en-n*c gypStUll deposing as those in the, upper Soudl-^Sast 
and near Menin^ir; (J ad 1921), and gyps Km in buned profiles near imudalecr 
( Stephens el a! 1 94 5 > where conditions a:v ;on humid for then Tun-iad-ii hCa> ; 
and fe also provided by fgg occv.rreure of fossil calcareous sod horizons under 
climatic conditions in which liixw pan acerivejation would not he expected to occur 
at present. Such fos^l II horizon > are to foe found in the "' travertin j' 1 ftl the 
tfordcrtowri district. In Western Australia r.rjionet/ soils occurring a: Rocky 
Gully, present aimtud rainfall over 30 inches, indicate. ffrOtitej: aridity {Stephens 
1946) than at pn-rcm. 

A considerate body of independent da a. much of which is of a penological 
nature, has hceu advanced therefore trj surest greater aridity in the past, and 
most of it appears "valid/' On the other hand, some of the evidence as the fact 
that the clunr s_\ 5;ems are now vvqelaml I Kiffs 1939. Crocker 1941 ) is of little 
or no value in itselL Kvideace of reduction of the effective area ru "active rvpc" 
dunes in south-western Queensland, tis advanced hy \V nuthouse (1940) "and 
neccpied hy Ihrovne (1945), is i&ki of doubtful va f ue, for the 'Vlead^ low, 
c6iup!e!eiy-vc^era'cd dunes v h : eh occur as <;mdu rs !o ;he Simpson De-err c':.n he 
matched by similar low vegetated dunes within the desert proper. While no 
suggestions of the present beint; ntaf* and than the period (ii maximum done 
huildin- have been advanced, it hu> been sfated that certain buried piotTes, ;ts 
those in the alluvium area? of the Noa.r'unga-Scl'jcks district of r South Australia, 
may iudinue greater pod.-olisatioti than the super-imposed present atfil and under- 
lying red soils ((cocker 1946). It is very diflkult 1o assess the age oi d:c.-e 
profiles, however. They could indicate, though ni; \\ fa uecessarv to 
proceed cautiously, tfei rl ere has been a yfrj rerenl decline following posi-atdd 
wetter conditions. Wbitehoucc (1940) tends to such a conclusion on die 
abundance 01 aboriginal artifacts in the far wesi of Queensland, m regions where 
there is now no permanent water. 

While further evidence is still desirable on some aspects, it wouid seem that 
( 1 ) preceding the onset of arulny ei'mruic conditions were much more luuuid than 
at present; (2) subsequent to the maximum aridity there was an increase in ram- 
fall, though not of a high order, and indeed very sired! by comparison with the 
preceding th-ererise — otherwise gypsum and h'nic accumulations, of the drier jveriivd 
would have been entirely removed in solution; (3) there may have been a vcrv 
n-cent minor decline irj rainfall, but considerably mure paleoper]o!ogicEil research 
will be needed to further elucidate this, in short, climatic conditions todav do not 
(idi'ei- greatly rfc-nt those when dimedmildiug was ai iis iua-:niium, but rain tall is 
;ippartntiy slightly higher. 

Tite d'scussion so far lias been concerned chiefly with rainfall as an index- 
Mi" ehmatc. Temperature e.Tects are much more difficult io defrrre. it is apparent- 
that It*" temperatures of the late Plcb.toeere, which coincided with ihe last preat 
glaciation, were low, so that in addition in higft rainfal) jirecediny the initiation 
of aridity, we can deduce colder conditions. 

The extinction in southern Australia of a suite of surface-dwenbur marine 
species dominated by dnadaru inrpczhi, which arc now tu be found \\ mam- 
locaTities alon;^ the southern coast a^ sidj-fossils. is of particular interest. TluVe 
specif rce still livir^ in more ttdrjJn»Tn \ Warme-;) waters, and it is considered 
their \rry recent cxtinctitai \\:is ( ;u.- nn a slid leu decline in temperature. It i* 


also certain that the suite, was present in .southern watery for a very limited time. 
The ^ggestitJn has been made that the brief stay of these species was coincidental 
with arid consequent upon the high Lenvpcratures associated wftb aridity (Crocker 
1946). It has not been suggested at what stage in the development of the desicca- 
tion Anadara and its associates became established along the southern Australian 
coast, hut it is likely that they persisted beyond the period of maximum dune- 
building (Crocker and Cotton 1946). 

There is strong evidence, therefore, to indicate that higher temperatures than 
those prevailing at present occurred in South Australia a very short lime ago, and 
these were probably coincident with ihe lower rainfall of the arid period. 

Fi^. 3 

T ic recorded distribution of dune sysuMiis and dune sheets, in Australia, 

(Chiefly after Madigan : HiflSj Prescoti: and Crocker.) 

3. Gknkral Effects of Akidity 

Loss of soil stability in South Australia and sun minding regions on a 
^rand scale resulted from the- ;-,nddcn modification of c!;ma:e. and it led to the 
build-up of the extensive .systems of aeoHati deposits, dunes, dune sheets, etc., of 
most of oiu: major and lesser ergs (fig. 3), Such Widespread wind erosion could 
only have occurred following the wholesale destrrclinn of the ioca! flora. It is 
apparent that for this to have happened the aridity must have been not cmly 
extrcmely severe, but its onset must have been pariieiilarly sudden. Orberwise 
the vegetation would have been able to maintain .so:! stability by sdrnple nitration. 
It was ohvionsly unable to do this. 

Jt is interesting, to picture whar probably happened. A catastrophic'' 
decline in rainfall which initiated tire aridity placed such l sTes.- on the pre- 
arid flora that over the greater part of the Stale and beyond it was almost com- 
pletely wiped out. An early replacement of rhis old flora bv a xeromorphic one 
capable of withstanding the aridity, thai is capah'e of million and ecesis imder 
Ihe extreme conditions, was not possible. This was chie'ly because the mo,st 

W From the biological viewpoint. 


desiccated region today, as for example, the Lake Eyre Basin, were, prior to the 
onset of aridity, so moist that centres for the rapid "dispersal of the diaspora of 
the required arid element mast have been extremely few, and perhaps almost non- 
existent. As the remnants of the more humid flora disappeared and disintegrated 
under the extreme desiccation, both wind and water erosion were greatly 
accelerated. The eroding power of occasional heavy rain?, which no doubt still 
occurred, but with probably greatly reduced frequency, was very high and carried 
considerable quantities, of silt and sand into ihc drainage basins. This, in the 
absence of a protective vegetative cover, -was transported and winnowed by the 
winds to initiate dune systems. In other parts the Quviatile agencies played a 
sma^or and smaller parr, until finally, at the other extreme, dune systems" of a 
purely deflationary origin on which iluviatile agencies were unimportant became 
piled up by wind (Crocker 1946) playing on soils which had completely lost their 
stability. This loss of soil stability was not general (see fig. 4), nor did it proceed 
lo Lbe ^nue degree or with the same 
rapidity everywhere. Certain soils 
wen- much more liable to complete loss 
m stability than others, depending, no 
doubt, upon a complex of factors, Tn 
£) >uih Australia soils which were 
derived from highly calcareous parent 
mali'tiil or which contained large 
qiuniluies of lime in IUq profile were 
particularly unstable. As a result the 
A hori::ons of the soils developed QB 
ihe '.it.: P'eislocene calcareous dunes, 
those HI our present malice regions, 
v, 1ih!i had received large quantities of 
calcium n< loess, and those derived 
from calcareous rocks, as the Tertiary 
Jime-'Tones of the NulJarbor Plains, 
were largely stripped, transported and 
re-soncd to form fcfgs. Typical 
ex;mrp'es arc the Nullarbor- Spencer 
dune sv=tem, and those of Ific South- PrjVtior,? °* South Australia where loss of soil 

! m and "mallee" regions. suUihiy *** l liht " f litTil j ° r . $m# during 

i, -.I., - , [ be imrJ-K^rent aridity. 

H is po>siMe mat strong wunJs 

greatly assisted the duneTmildmg, hm sum a possibility mav be dismissed for 

wani of any evidence one way or the other. Although nothing can be said of the 

intensity of arid winds, the general wind pattern was apparently very similar 

to that today, because the done systems throughout Australia, which arc entirely 

or almost entirely of the satidridge or self lype, show a consistent orientation with 

the whid regimes of the present ( Madigan 19<3p, Hills 1939). 

One consequence of the fact that" soils varied in their erodibihty and I hat 

possibilities for the maintenance of vegetative cover, despite the adverse climatic 

trends, varied in different localities (see later), wa, that the dunes winx not alJ 

initiated at the same time. For example, although resultant from the same arid 

period, the time of dune-building initiation and maxima mav not have coincided 

exactly in different localities. The Murray inallee systems may have been slightly 

in advance or behind those, say, fa the lower South -East. 


Although over the greater pafL of South Australia and the adjoining hinter- 
land regions the old vegetation was largely destroyed and countless species mu^t 


have been entirely extinguished, remnants of the pre-arid ilora managed Lo survive 
the desiccation in especially favoured situations. 

The pre-arid topography was almost ideurical with that of today, except for 
the purely superficial, sand deposits, It is obvious, therefore, that the rainfall 
gradients were similarly zoned. This pattern is determined chiefly by latitude 
and distance from the coast, but is modified by other factors, most important of 
which are the influences of mounUiin ranges and hills. It is apparent that when 
the severe climate stresses associated with the sudden onset of desiccation were 
imposed, the mesic flora's sole defence was a migration towards wetter conditions. 

Fig. 5 
Principal refuges during the Great Australian Arid Period, 

Successful migrations would only be possible where the rainfall gradients 
were steep — that is where the distances between climaiic horizons were small and 
where the propagule dispersion capacity, the establishment capacity and the ecesis 
capacity of the species were rapid enough. 

Reference to the present rainfall distribution, and comparison with a topo- 
graphic map, indicate quite clearly that the possibilities for successful migralion 
were very limited and could occur only adjacent to the ma; or ranges. The ranges 
were important regions of survival for another reason even where their influence 
on rainfall was not so great, for within them occurred the greatest diversity of 
microclimates. In the ranges and hills were the largest number of ecological 


niches, the greatest variation in habitats, to be offered the retreating vegetation. 
They were, therefore, the principal refuses of the rrlic flora which survived the 
stress tyf aridity. Other refuges of less importance were the major drainage lines, 
any of the lakes which, although reduced in area, did not entirely dry up, and 
oLher mitres more humid than surrounding areas. 

Tbc principal refuge? of the arid period m South Australia and adjacent 
regfofljs are shown cm the accompanying map (rig. 5). The tiffltytf imnortant were, 
no doubt, those of the Mount Lofty-Flinders Range system. Others of note 
were Kangaroo Island, the ranges of eastern Eyre Peninsula, the Gawler Ranges, 
the Mrr^raves, the Macdnnnoll-James Range system (Central Ausl), the Gram- 
pians (Vict.) and the Murray drainage system. Although these were the main 
centres of survival, the vegetative cover within them was very restricted. This is 
evidenced by the presence of climbing dunes in the Miitalie region of Eyre 
PcninsnW. These (Junes &**? outliers of. but more or less continuous with, those 
of the VuHarbor-Spencer dun* 1 system, and cover portion of the ranges of 
central County fervors. Similar evidence is provided by the aeolian sands 
of portion of the Mount Lofty Rantc system, as for example, those that are sq 
conspicuous in parts of the P>remer Valley. 


The ntimerotts ecological niches which acted as centres of survival were also 
the roe;il points from which commenced the migrations leading to the establish- 
ment of the present-day communities. Xo doubt the climatic duress associated 
with the defecation resulted in \nxt\t modifications in the flora. Tn all likelihood 
the isolation and continued stress of existing under conditions a;ioroa^hin.jr or 
at the, lunits of tolerance for most species, favoured a large number of sub-species 
and strains relatively unimportant prcvionslv, and resulted in endcmisni of varying 
decrees. It is interesting to trace the major aspects of this development and to 
Attempt a definition of some of the migratory routes and the principles involved. 


A'Htourh tfafl *rverify of the atid:l.v re*n!k-d m such a restriction in arm 
nf species r.nsl the destruction of I he* ore-arid vegetation on a grand =cnh\ it is 
prohrdi'e that a slow recoloni-aHon was be/run be n fen* drought resistant and 
perhaps some new genotypes, before Juty improvement in rainfall look place. Any 
snr't mie/nnion, however, nvi^t ha\c becri slow and limited to a few prcv^m-U 
uuinniortant snecie- with noor iltrift/^il capacities, for otherwise jfrfc In** of soil 
stfttiJIftv frlfetlft not have been pf» v\imfr\\ti£zH or have prorec Igd so (Ufa If \* 
obvious (hat the greatest nnmbev of spec* : es snrvivhie in the relatively few refu£f> 
were far from bcimr under opthnoru condition-, aud competition f(ir the available 
ecological habitats was grctyfe H was v/>t until an improvement m climatic con- 
ditions, which probably included an increased rainfall and r-bVnfV lower :em- 
perafurcs, resubed in a release of both the pressure of competition and the uk- t.vf an adverse ehmate that r^-rnVm^.rion coidd b? very eiT?ciive. 

The sp^ed at which r«- enioiu^rdion proceeded ;md tile pattern it followed 
wou'd depend on many factor-:. In t'.c tdtimme r.'ialv-ds i: ifrflrrH htf eovcrn-d 
by tb> 1 abilily of the individual supcrep artrf i's biotypes to produce and disseminate 
pror«r,'?u!^s. The speed with \yfiitjH species could exploit, the npporrunilies of Hi 
e-t. nnV;1 climatic habitat ran^e w<mH depend, therefore, on their especial 
mecr-amsms t'tr dispersal. Abhniurh mo^l p^ant geographers believe thai Ihrtg 
distance dispersal is of infrequent occurrence owing; to the effective barrier of 
< Mablished communities, such a deterrent would not be opera' ive in tltis case 
hecnuse the area hennr coloniser] was virlna'iv a bare one. ft is ltfcelv, therefore, 


that with the relaxation of climatic stress the species with capacity for wide 
dispersal of jrermules rapidly became widespread and, especially away from thfc 
centre* of survival, played the dominant role in the early stages of the sere.i.. 

2. Migratory "Routes 

Dissemination of propagiiles is, of course, only the first stage in the process 
of re-colonisatinn. It is. necessary for them to germinate and become established 
in the new sites. Only germules which come, to rest in suitable habitats, that is 
habitat within the ran^e of tolerance of the eeotvpc, can hope to survive and 
eccsise. Within the climatic amplitude of the biolypes, atec£Stf fill establishment 
of species in the invading flora would be governed, therefore^ principally by 
oibrr environmental factors. In the relative absence of couipethion in the initial 
stages, tbere ean he little doubt that the distribution of die early elements was 
determined whhin iheir climatic tolerances, chiefly by Hie 1 edaphie environment. 
Soil condition* over and above the effect of chance distributing of die initial 
proprieties, were undoubtedly die major ecological agents which jnndin>d the 
checkerboard of the invading vegetation, and indeed, imposed upon it a special 

Although the under! vine; causes and the modes of dispersal are fundamental 
to distribution, they would have had little influence in just where Hie propaiiulcs 
initially came to rest. This was dependent upon the relationship of the parent 
plant to the agents of di?per?ion. physiography, etc. 11 rt is, chiefly dependent 
upon meteorological factors, waterways and physiographic condition?. The exact 
site nl which a propa^ule lodges and germinates is, as has been pointed out by 
Cain (l fl 44), entirely clue 10 chance in the sen^e tint tliere ;s no conscious 
co-operation between the gcminle and the complex of dispersal agencies. Follow- 
ing dispersal, ^emiination. and m suitable habitats ecesis, aggregation and re- 
di^persal of species no doubt occurred, and very soon the factor of competition 
became opera'ive, imposing a general, diou^h variable, brake noon the rate- of 
migration. Nearer the centres of survival the harrier o£ competition would 
obviously he imposed earlier because of the greater density of propaguies. 

With the release of ch'matie pressure on the relic i'ora, re-coVwnssrin'i, no 
cYmht, commenced from most, It is apparent, therefore, that potential 
inigrnlion could have proceeded in many directions a: once. The. extent and speed 
at whitli these movements took place was governed, apart from thgsc t-i.ctors, 
reside m* widdn the specie^ itself lilce dispersal capacity, rlr'efly by the soil m.v-aie, 
a- mentioned previously. Tins is. indeed a pi iiiciple of dynamic plant geography 
(Gleasnn 10ZV). It has been stressr-d hy Cain (1944) in sayinp; that sueces^frd 
migration depends upon the occurrence of a "continuity of habitats winch are 
within th'd ecological ampln'tide cd |}r* i nip raring sp;:e:c*s." Tbe "ecological 
jmrplio-de,' 1 of a species lias hs scat in if. genetic eonslhmdou, and strictly, there 
fore, vuthfn the species itself, but tl:e occurrence of stihnhle habitats within this 
ranr^c is chiefly a function of exoiojx:-; id and pedo:o;;iad history in which w/'-ta- 
t'on plays an important, but minor, rck\ It is almost cc.oun, therefore, :hat lite 
main moratory routes in the deve!o[ mart of die present Stfutji Australian dor a 
have fxrn from die iun-dvaf cciUree. atttt^ series of e'nsely r/afcj S&JJs, 

The extensive dune systems, many of w1. : en are Imbed directly w»i i £lie 
survival centres the Spencer- Vmccnf -ysleni. Murray ^lnllt-e dunes, ^nnn-on 
Desert, etc. — would, for example, have been admirable routes foi re-colonisrtion. 
and were, no doubt, some of the mast important migratory tracts. Gibers were 
the gibber downs, the mallee- de-cri ioam soil complex, and drainage line:?, both 
minor -and major. The ma'lcr-oesert loam soil comples. coves a multku de of 
variation, hot w.thin d there h a repet irinn n" closely ndaied fdaphir li.-er.jats 

which are raaf beyond the dispersal ratine of many species. It vva=> quite clearly 
out oL ilie major migratory Indeed, in the broad scuse, and omitting the 
draitra-e lines, the aeolirm Hand systems, the gibber downs, and the mallec-desert 
:oam <uil complex have I&CU the three major edaphie influences in the develop- 
ment and distribution of the present dora. To these may be added the red-brown 
irfevl.%1 which extend in ft practically unbroken sequence iTom the A«)elaide region 
to t\v: lower parts of the northern Klinders, and the terra and tlieir allies 
Hrbicll are associated with acolianite limestone. 

fia addition to fcbe importance of series of closely rolaied e.d.iplre. habitats 
wi'bin die ecological tolerances of the advancing -peries, climate has been a dv 
l' ■tmm.Ui/ Influence on the direction of the main highways. As ihe ini^ra'.ioir; 
re, \> I their greatest impetus with a small increase in rainfall, it LS obvious dial 
liiev \f\'i tceeded progressively (Vop1 more humid to dri^r areas. This is well 
iilrsii.i'.ed by an analysis of the principal elements of the vegetation of the lower 
Sonlb-r.asi ol' South Australia. Of a total number of 212 Australian Species 
recoi\!c:l on a general ecological survey of the region (Crocker 1944), 136 were 
limited to t:;us:crn Australia and South Australia, 52 were to Ifci found in both 
ea<t ano 1 west Australia, n 1 1 T 23 were restricted to South Australia. One. species 
(a g?&£&) was limited to South Australia and Western Australia. It is obvious 
that the major re-coIonis*atinn in this region has proceeded from the east, that is 
from survival centres in western Victoria. The above analysis was made on the 
vegetation as a itf|jol<2. A separate analysis of the ■sclcrophyll communities, which 
occur on sails that were completely unstable and therefore devoid of vegetation 
during the aridity, would httVti shown this trend even more strongly. 
for it is most likely that a few isolated swamps of the in tor- range plains of the 
region did noi become completely dry. They probably acted as focal points lor 
re-diAiiibution of some species like Gchnia h'ifuia, Pna australis, etc. 

The principal moratory routes in the. re-establishmem of vegetation cover 
and the development of the present communities in South Australia art? shown 
in figi f>. They have been superimposed upon a somewhat simplified soil map 
bas'cd on i.hat published by Prescort (1944), together with information collected 
by one pf the authors (1\< [* C). The interpretation has beeu modified to suit 
the requirements of the present paper and no attempt has been made to indicate 
thu migration of types, such us the occurrence of malice or gibber soils in inter- 
dune tf uriclors, etc. More detailed information on soil distribution in South 
Australia can be obtained from other publications ( Prescott 1931. 1914, etc.). 


The problems of ecology*, and in its broader aspects plant geography, are 
laryivy those of explaining the types of species distribution and the differences 
bet neon plant communities. Because species and their biotype.s react differently 
to changes in environmental conditions, that is, because their potential cdapho- 
efiflinlir environments vary, in the uirmatc analysis their distribution is governed 
by tht- Individual ranges of tolerance. The individual may ? therefore, have a 
pMicntially wide or narrow distribution, depending upon the. occurrence of suit- 
able e'miaiie and cdaplue habitats. 

If has been difficult to ohuuu an independent as c essmcnt of tile effect ot 
strife and climalc upon the distribution of species within an association, because 
ebuiai'- pia>s a dominant role in pedogenesis in determining both the ^cochemicat 
trends and the morphology of the limd soil profile. The matter jc tnriher com- 
plicated hy the problem of possib!e migrations and their direction. A unique 
upponunity wa* t nVrcd to study the response to changes in climate in the lowrr 
South-Exist of South Australia, where both the* direction of migration was known* 


and was predominantly unidirectional, and a series of closely allied and largely 
identical soil types extended, because o£ an unusual history, across the climatic 
zones, passing through several of them. The effect oi climate was seen to be a 
gradual modification of both specific composition and structure of the community 
— a Eucalyptus Baxtcri sclerophy'dons forcsh As species approached the limits 

' ir/o ii-« « i ■ » I 

* ,' - '• • • • •_ i: 

PocLsois -Qsid. Red- brown, -eant&s 

T&rra. ro$&CL - RcrLcLzi/ux. cuff* 


Fig. 6 
The principal migratory mutes. 

of their climatic range on this soil they became dwarfed and depauperate and 
finally dropped out (Crocker 1944). This "sifting" effect of climate was first 
noticed by Good (1937) in arid communities. On the other hand, soil changes 
usually depend on geological history, and vegetational responses to them at the 
same climatic horizon are sharp and clear-cut and are by no means difficult to 

The results of the studies cited in the previous paragraph led one of the 
authors (R. L. C.) to the conclusion that *'edaphic factors .... are responsible 


for the distribution of formations and associations within any climatic horizon " 
While this generalisation was, in the light of future experience, only a part truth, 
it could be made quite accurate by the addition of the words, "other things being 
equal." In practice, except in restricted areas, other things are not equal. First 
and foremost there are varying migration capacities and the possibility of opposing 
migrations. For example, it is unquestionable that communities dominated by 
species dispersing eastwards from the Mount Lofty-Flinders Range refuges met 
somewhere the barrier of vegetation migrating westwards from easlcrn centres 
of dispersal, such a? the east coast mountains or the Barrier Ranges, and vice- 
versa. Migrations in all directions was obviously going on simultaneously and 
influenced the distribution of communities. It is better T however, to bring the 
discussion back to the species level. While some species within a community arc 


Fig. 7 
The recorded distribution 01 bladder sahbush, Atriplex vcsic&riutu. 

dependent in that their habitat tolerances require the shade or protection, etc., of 
another of a higher stratum, an understanding oF the factors affecting the area 
of non-dependent (chiefly dominant) species will usually provide the key to an 
understanding of the former. 

Within narrow hunts the chance settling of prupagules determines the exact 
distribution as pointed out by Fidgeon and Ashby (1942). In the broader sense, 
however, there occur many other interesting types of distribution which cannot 
be attributed entirely to chance. Firstly, rhere arc distributions where over wide 
areas within the one climatic fcofleW the same species keep recurring. Typical 
examples are the malices Rucalypius oleosa and E. gniciliSj and saltbush Airiplex 

( 2 ) Climatic zone- i$ here used 
vidual species. 


rise of the zone of tolei 

for tht: indi- 


vesicarium. .Secondly there is the tare of distrihuti'n, and one that is more 
puzzling, of a aperies like Arociu feme (vvaddy'l, which occurs as two small 
groups of a tew hundred or ks.s trees on Lh$ ^thber downs, now separaved by 
approximately 200 miles of Simpson Desert sand (Crocker I°4(V}_ Another type 
of discontinuity is that of the sugar gum ( Eucalyptus cfutirnuixx), which occurs 
in ihrrc relatively isolated rcginfK in South Australia, namely, restricted parts of 
Kangaroo Island, Jvyre Peninsula znd the Southern Flinders Ranges. Can? 
(I04T). in his valuable thesis of plan; geography, has applied the term disjn;wtit>n 
to tho^e distributions where inrlividual species are separated by more than the 
norma! dispersal capacity of the ivpe, and concludes that major disjunctions have 
almost exclusively resulted from historical causes. 

1 1 is obvious that the development of the South Australian flora which has 
afrendy been traced in outline, has been such that -ceo lo^Lal and plant geographical 
pvnbVms in area! distribution of species will only be understood clearly if the 
historical background is appreciated. At p:esent the restricted knowledge of 
pa!*.*' botany, pa', and pa'eormuaioloiTy makes run thing like a full 
evalralmn impossible. Willi the start thai has already ircn made, however, it is 
worth while to raise some of the problems and anomalies and to attempt an 
explanation of the more striking discontinuities. 

1. Winr; (con'm.nuous) D:ftTun!UTio.NS 
The present recorded distribution of /ttr>plcx vexiciriniH fe interesting- It ifi 
shown in ihe accompanying tijjiire 7. Deat'hig in miud the principle thai dispersal 
proceeded chiefly from weLter to drier habitat*, it is likely that Alriplcx xcstaiyhtw 
had .-le.vernl centres of rcclislribui'on. tig isolated occurrence in the Maedonnell 
R;:nges suggests that one migration emanated from 'here. The northern Flinders 
arid Gawler Ranges were also probably centres of survival and redispersal. Nor 
is it necessary to believe Ilia! there were not o hers. Th* two uia!k*ee», Eucalyptus 
oleosa and Eucalyptus gracilis 1'kewise probably survived the aridity in geo- 
graphical uolat^on in several refuges (Eyre Peninsula, Mount Lofty Ranges, etc.). 
from which they have subsequently spread (>;ee fig. 8). The Victorian and 
Riverina occurrences are probably migrations from South Australia. Wide and 
continuous distributions of these types are merely indicative of the occurrence- of 
extensive edapho-clinmtic habitats within the tolerance tanges of the species, and 
an ability and opportunity for wide dispersal, for establishment and ececi.-.. The 
two malice- 1 : and Atriplex irswariuin occur on jfoj desert lonm mid tec soil 
complex. The two soil groups merge more or less gradually into each other 5i 
Sonii Australia. While the elinranc ranges of the two imdlees are less arid than 
those, of Alriphx vesicarlutn, which suggests more humid and probably more 
southerly centres of survival for them, the distributions do overlap, and towards 
the limits of their ranges the three species are frequently found together. AtHpl&d 
vcsiairiiau is a species with fairly wide edaphic tolerances. Tn addition to grow- 
ing on mailer soils and certain of the- desert lonms, »t is able 10 cstabksh and 
feesisc on some of the gibber desert coils and eertain rdtrvkd soils of the Rivcrina. 
Because of these wide tolerances it has not been difficult for it to find a series of 
suitable habitats within its normal dispersal capacity. 

2. Restricted J3iSTEirajxicxs- — Rri.ic and Exofmjc SrEClUK 
There are two ways of regarding species of very restricted distribution, either 
as young endemics or as old endemics, that is relics of a previous flora. Willis 
fl938, 1940), for example, considers that mo^t endemics are young species rather 
than relics. Ridley (1925) suggests the tctm rpibioiivx for rciic endemics. JVluch 
doubt and considerate difference of opiuioti still exist? about the ln*t r rrlation 


ships of the factors causing" speciaticm. but it lias nevertheless been clearly demon- 
strated that -isolation, as in mounLain systems and oceanic islands, results ta a high 
cteggcq of endemisin. 

This is well shown in southern Austra'ia by the high percentage of enclenrism 
on Kangaroo Island by comparison with the adjacent peninsulas (Wood, 1930). 
An analysis of the distribution patterns of the sclerophyil communities ill these 
peninsulas showed that of the 82 species endemic to the region as a whole, t3B$fc 
were found on Kangaroo Island, and 47% of them were restricted to it. The only 
other peninsula with endemic species was Eyre Peninsula, where the remaining 
12% occurred. Lt is perhaps even better demonstrated by the comparative 
endemism in southern Australian eucalypts (Herbert 1928). In the south-west 
corner of Western Australia (Shark's Bay-Eucla districts) 74*2% of the 


Fig. 8 

Tiro verore'eel tfistrii-.tition oi {Up two mallei TZite&ltphtf. oleoma and 
£.. gracilis m Australia. Their general areas are almost identical. 

eucalypts are endemic to the region, wlvch ha; had a long and undisturbed 
geo!og:caT hlsiorv. IVmpcratc east Australia, geologically a much more disturbed 
area, and connected by more or less, continuous mountain chains with the tropics, 
exhibits endeinism of onh' A-H-7 r /c among the cuca'ypts. The case for high 
endemism in geographica'dy isolated areas is. therefore, supported by Australian 

ft is oi considerable interest in assessing the status of species in our flora to 
know whether the endemics are young or relic. During the recent extreme aridity 
the climatic stress imposed a rigid selection upon the Australian plant populations. 
This, as pointed out earlier, meant the preservation of species in which the 
biotypical range was considerably modified. The overall result was the extinction 
of many biotypes (and species) and the preservation of others. The areal extent 
of the present derivative can for this reason alone, i.e., climatic selection, have 


[iuie relationship to the distribution of the pre arid parent stock. In addit on to 
the direct climatic filtering- to which t lie old vegetali-un was subjected, others 
equallv important in determining the present distributions have been super- 
imposed. First there was the effect 61 desiccation upon the sons themselves. Tins 
influence was profound, To varying degrees over mo^t of South Aysiralia sotis 
lost thejr stability, and, under the accelerated eroMon resuinng,_ were stripped of 
some or all of their A horizons which were re-sorted and piled into dune systems, 
nr spread out as sand sheers. Wheu ihc re-colonising Specif began Ld spread 
rapidly with the release of climatic pressure, they were coin routed then with an 
entirely new (in the narrow sense) or drastically modified on the broader sense) 
soil pattern. 

From a consideration of the foregoing, the conclusion seems to be iha: the 
present distributions, except where the climatic zones are i:.arrow f cannot no more 
than give one or two glimpses of tfir flora pi the pre arid part. 

An undoubted relic is Livlsfana Marine, and another Man'osomnt Mac- 
i!nnudi{, which are found in a single valley, the Glen of Palms, in the fiacre nnel 
Rondos, Central Australia. It was suggested by Spencer (1921) that it was a 

t en mart of an older flora and 

; indicative of a wetier ehmatc 

in the pasL There seems no 
other explanation that ;ott1d 
be applied. 




appears 1o be relic is Acacia 
prrrc. Its known distribution 
has bout already descriijed and 
is shown in fig 1 ; 9. Jt 's diffi- 
enlt to imagine that k :oukl 
be a. » 1 e w speci cs , i or the 
w :y \ mi i etj occurrences are 
separated by an unfavourable 
edaphie habitat, namely. 200 
m'les tii desert sands. The 

The distriontion of \vaddv\ Acacia pats?- This h& 
spcctacul.-i!* tree gtbtfbg io 5U feet high. 


is far beyo*i:.. the 
capacity of the 

s; -ec es. unk ss long dis- 
o.see d ; spersrd had operated. 
Allhou.h di>pcrsal hy ageeis *ueh as birds (Ridley ' ( W) can occasionally 
be responsible for JWiwSe disjunction?, it is not considered likely to have 
operated in ilvs case. For otie reason then: nppear Ic hfi ample ivnoceupicd 
habitats within the edapho-climalic amplitude of the s penct. <md l( ir >vW= 
capable of dispersal, estabiishrr.^rii and ftttus over- a ninge of 200 mil:^, it 
seems extraordinary that it has not made uvre progress nearer home, even assum- 
ing on:- or two occurrences have not yet been recorded. The. most likely explana- 
tion would appear to he that it :s a true relic, which Avas formerly widespread bul 
was restricted to ecological niches in widely separated localities with, the onset 
of aridity, louring the desiccation the desert sands piled up between. With sub- 
seouenl amelioration the species has migrated and spread, but on'!}- to a very small 
extent: perhaps n is still inwards the limits ot its lo!erin:es. 

Ouc gt the mo-s imere^mg cases nt d^couiinmiy in S^iith Australia is * ;&% tjit 
sugar gum. Eucalyptus cladocalyx. It occurs in three widely spaced localities 
(see fig. 10^ : (1) Southern Eyre Peninsula, (2) Kangaroo Island, and (3) the 


binders Ranees. On Eyre Peninsula it grows chiefly on t! 






residuaU and podsols, ainj is panieularly depauperate. It has been suggested 
that it is towards the limits flf its edaphic-climatic range (Crocker 1946). On 
Kangaroo Island it flanks creeks and nrrurs on soil a.ssoeiaied Aviiii the older 
sedimenraries. Tt is a much more handsome tree than on the adjacent Eyre 
Peninsula. The lower Flinders distribution in limited to the. neighbourhood o[ 
YYirrabara, although there arc several outliers further north. 

In oi;r opinion the three areas of Jlucalyp'tts dadocutyx have developed from 
three centres of survival, and are mu j reuc?. Why the specter has not spread 
more is dirrini'it to understand. tecauR: its ttS« m homestead plantations 
that it. ha:v a wide cdaphic ran^e. For Home reason the specie--; has a poor migra- 
tion {pU-te establishment) capacity. In this connection it is necessary 10 remember 
that r an:jen of tolerance, that is minimum, optimum and maximum levels of 
tolerance to any ccologjc factor, vary fft different stages be development. The 
possibihty that Eitorfyfivs ctadccuty.r H a new species and that its present dis- 
tribution patlern is due to king; distance d^pcrsal, j s very difttcu'i to believe. 

T*. . :■ : ;i+erestiug cases of discoiiriuitity 
l.-iyij i'_tru observed recently by C. I). 
i\in ....;•.,-;-.. One is the tedisc'A cry of an 
a*^e. r.t rM siviu^ybark, jitwuivpttti 
ir-:,\ -:■■'': y;:cfra, near C ! arc\ South Ails- 
* ' -",[ : :.'e nce.iv-.t recorded occurrence 
is :"1 T^'i Grampians ox Virlona, The 
other Is xhc occurrence of grey ironhnrk, 
/: .s,.'; 1 .: /.<• lK r i)i:;>h!oiu, in the northern 
l : U:\\v> Kanges; ;he nearest record is 
i , a i a-e occurj ence i.ear ,! ! orsham 

Ahhoiu;h ihc general apnucaCon ot 
Hiii^ distance dispersal at a means of 
evp^ri -'nig discontinuities has been largely 
( : ^;:v '-.^4. I.jy the work of SkoLt:d;crg 
ib' : ?. 1^0 )' r.nd o:hevs, the possibility 
of ;i» r.pjraiiug in specta 1 cases has been 
r,i' :ly u-Jmiuc.d, The- urn:;; argume.his 
n t '•:■; :)'c theory have res:ei upon lh* 
p:\ vtu distribution of ctidcm'cs in relu- 
oou ifi ay of halation, an 1 because ox 

the diffkuhy of demonstrating that far travel'ed germuks could cuter and compete 
in the communities already present. In the early stages of the re-co-onisarion of 
South Australia this competition factor would be teas important, because, as 
pointed out earlier, the area was virtually a hare one and the chances of successful 
long-distance dispersal must have been greatly increased. Such conditions would 
be short-lived, however. Long-di$tr,nce di-per.sal is undoubtedly accomplished in 
the ease of Hght gcrmules; like the spores of mosses and lichens (Rirriey W3Q, 
Baas Eeeking J9o4). These, in addition to being widespread, frequently develop 
and grow in such extreme and selective environments thai the problem of corn- 
pet-iion in an already closed community is less frequently operative. A case of 
distribitoou of this type in .South Australia is that of Sphr/ywim, which is found in 
two or three small swamps near Mount Burr (Crocker and luirdley 1929). The 
nearesi recorded occurrence of Sha^knum is near Melbourne, approximately 
250 miles awa>. The more e:.-m-iur a id specific ihc environment, Ihc fcss likely 
is* competition, and the more likely is long distance dispersal to he. operative. 
CUt&s-ie examples arc the distribution of ${*ffpi# m^rithna au,d the y( Dumdiehoii" 
community (Wood and Uaas Becking 1937). 

CmoJcib-IUs c£adocaiL-.K 

Ftp 10 

The natural distribution of Sugar j?mn, 
£ti<a!yp(us cladoca'yx. hi South Australia. 


A special type of discontinuity is that of the two acacias, A. Sowttttiii Maiden 
and A. Ladcri Maiden. The former is conspicuous in the Tarconla-Bucklebuu- 
Port Augusta area at the north nf Ryre 3?enuto% while the latter is of limited 
occurrence south of Cockhurn along the Ne.w South Wales border and extends 
into that Stale (see fijr. 11). The two species are very closely related. Thev are 
almost idcruVnl in habit, appearance mid other morphological characters. They 
occur within the same climatic /one, on wry similar soil l -/pes, and art: associated 
in ilir comimuihies they dominate with a targe number of ihe same species They 
differ iavoromieally rhfr-fly in the length of "the phylkxna (4-8 cm. and 7-11 cm. 
respectively), and in ihe pod o\ A. J.odvri bring mor.iiifonn, while that of 
A.SmaJcnii is but slightly constricted. Species of this lype which are only sightly 
different morphologically, but which are geogr:- phi rally very isolated, hare Wn 
defined ft? pt&m&t* (Selcbell 19,35). In the author's opinion thc'abnvc 1uo 
species have descended from ihe same parent species, once widespread, and are 

merely the product 
of bie-tvpienl isolation 
within (hat species. 
It tfi likely that the 
(osirihulion of Acacia 
$Oiyd\ 'Ht'r, for example, 
has developed either 
ironi a survival centre 
in the northern Flin- 
ders, or, as is more 
likely, from ihe 
(iawler Ranges. On 
the other hand, Acacia 
i -"iter, was distributed 
from .some oilier re- 
fuges, probably in 
Mew South Wales. 
Under the differing 
cdapho-ehmaiic sclec- 
minor characteristics 

The (Yujtmcttr.n between the vicari'mis species Acacia Soivdemi and 
Acacia Lodcri. 


turn pressures in the two regions biotvpes which vary 
have been preserved and have been given specific rank. 

Depending on the point of view, .4. So-wdcuii and A. Lodcri can be con- 
sidered either relic (old) or yoitn^ endemics, for obviously they arc both. The 
relationship between age of specie.-; and area as proposed tiy Willis (1922) would 
relegate them to the position of young species, on the other hand they are certainly 
reiic in that they are descended from a previously more widespread parent that 
was restricted in area as a result of unfavourable climatic development, Even 
better evidence is presented for relic slulus by Ihe distribution of species like 
Livisiona Manac and Acacia fmG& In cases |0^ these the historical sequences 
indicate a reasonably good hasts for consideration as relics. 

If is essential, however, to retain the dynamic concept of vegetation and to 
realize lhat the historical changes, which cause contraction tn area, may preserve 
in some of the old populations strain eha-acteri sties that wil! allow rapid'expansion 
of area for the new edapho-elimatic conditions when the restrictive pressures are 
removed. In other w orris, relic species can be widespread, as well as confined in 
area (epibiotics j. In the broad sense, therefore, species like Afyiplcx iedcmtw 
and Lucalyptns oleosa may be. and probably arc, true prc-arid relics. Voting 
endemics might also have become widespread in South Australia, provided their 
ecological amplitudes were wide enough to permit dispersal, establishment and 


ecesis either on a high level ot competition, or if they had developed prior to, or 
upon Hie release ot climatic presume, at a lower level of competition. 

The Acacia Sowdcnii and ,-1 Lovrvi discontinuity has been considered^ an 
cxamuio of biotvy- isolation of a more widespread parent specie? or population. 
In vi«-\v of the new historical data ir would seem that some of the cicely related 
pairs of species (Wood and Baas Becking 19.57), j n the Eucalypi seierophyll 
iore^ ot the Brtie Mountains region of Kew South Wales and to '.he Aloun 1 
Lofty Karnes. Soulh Australia, might be considered to have had & similar ori^m. 
It is not unlikelv tliat in the pluvial pr-Arid. that is, in the lale PkaVoeenc and/or 
the carlv Reeenl. or at an even earlier tim?. a series of suitable odapiio-ciimanc 
habdals'odsU-d IQ foid^c die intervening; area. Isolation followed with the aridity 
(or earlier), and has rented in differential biotypc selection. *\lau> of the pans 
therefore, arc probably related historically to a common parent, and toe an examp.e 
of divergence rather than convergence. 


Tiruofeef-"Rcs*;ov5ky [V)A2) has applied the reiui ini<:ro-ei'o!v,iii>h to vai^tm-i 
for which scientific evidence of mode frf operation can be obtaiuc L Specia;;on 
and geographical differentiation belong to tfe class. As a result of much gene-tic 
work considerable agreement has been reached on the factors involved in micro - 
cvolutioii, and it is agreed thai mulauon is the mrdn mechanism for providm- 
new evolutionary material. Cytogenetic work has also shown that fin 
specinriou is ovnerally due to the ;icaunuhi( : im of small character difference-, 
and no; to giftgte malarious as claimed by Willis ( 1940). What bttle is known of 
tnulation rates $ttgge&$s that thev are Of a very PttV order and that they arise at 
random ( Sewed Wright 1940 I. ' Evolution, or successful differentiation fettiMng 
in speciation is the result of factors n:hci than imitation shaping the evohuiona;*. 
materia!, ami of these selection and feofyltou are most important (Timofeei- 
h!rs*>vJ:y i C), ff). Dariimdon 1910. etc.). The role of the different factor* in 
evolution no doubt varies under different circumstances as pointed out h\ 
Diver (J9d0"l, nor is (here. ;iny theoretical reason for believing" this is not so 
(Sewen W*5g*tt 1^-0). 

In iJbp foregoing sections realms have been piven for as-uuiing a large prc- 
arki relic elc-iucnt in on r flam. Cert;dn discontinuous di-trihutiuns have hefu dis 
cussed from tine ftngie, because ;i vthV 0FJg*Jl av^e^ars to he the mo.-.t logicaJ and 
straightforward emanation of ibeir pre- fi1 aie,c-. 1: is obvious, however, that 
considerable revolutionary differentiation ftrta prtitert% occurred since iIk om<t 
of aridity, some of which wnu'd have resulted in full speriatioit. Further, t\W 
i-plaiionYesuUiug from the eomraclion of the liora must have favoured specKi(:on. 

T-o'ation is considered a directive factor in evolution by Timofe;ddU\ssovsky. 
The chief importance of geographical isolation lies in die restriction of free inter- 
brcedincr, yvhtck rrsulrs from partial isolation f>f a population, f'ecausc of thK 
isolated Tortious ol the original population vvhh differing evolutionary potentials 
plight giyK me, to nmUuious which come under differ-san selective pre^mc-s. and 
final lv result m considerable divergence within the different centres of isolation. 
The ab-orptton ot" differential ions resulting' in tins way is probably prevented by 
the restrictions to interlu-oeding-. tl is possible that in this manner, or something 
closely L'khi, repro- active isolation and hence full speciation can be uhhnately 
achieved. Chi the oilier hand, reproductive i^olai'on tuay be more frequently tbe 
TCSttlt of hyliridis.aiion and intercrossaig following the reunion nf adaptive com- 
plexes developed under geographical or ecological isolation (Dhobzban^ky 1941). 
In ddier ca?e geographical i^ilaiion las played un Inipovlant role. So far .us fljc 
V\uslmlian flora is concerned, the contraction in area associated -with them id 


period mast have been an important ditecivc factor promoting- biotyptcal differen- 
tiation, b'urther, on the release of climatic pressure, the rapid expansion of urea 
of those hiolypeji able !o exploh the new edapho-r'imane habtlats mini have giv'tm 
some excellent opportunities for reunion — dial is i*or hybridisation, baelc-cros> "ng, 
and various other types of imercrnssmg. 

It hvjjHt be expected then, in view of the history of the. Australian flora, and 
the present state of knowledge of factors effecting spec:ai ; ou. thut considerable 
differentiation ha", occurred since the isolation of the Sorn in numerous centres 
of survival during the Recent dc -dec-atinn. \Iauy of ire: e divergences; have pro- 
bably revolted hi full speciatiou. One limiting far or i^ undoubtedly the relatively 
short lime involved. liven assuming that Hie arid period was centred much longer 
ago than i.> at present believed (4,000-6.01.10 year?), it \va^ probably not more than 
10.000 years ago. Considering the g:-neraliv low order of dose mulaiion rah's !: ' J 
which have been measured by genetici-Sl.s, and the Filet that the grent majority of 
these in any case have no adaptive value, the amount of [ml specu^ion over 'his 
period must have been limited. It would of course depend, amongst uLher things, 
upon the degree of both potential and developed diiferen.atiuu <n a particttbir 
group pri.or to contraction, and this would have varied widely within the differing 
species complexes. 

Geographical and ecological isolation are very closely related and grade into 
each other. Tlie importance of ecological isolation in differentiation, however, 
does not seem to have been sufficiently stressor]. Wkh vcgeiation, area! limitations 
within the climate tolerances of a species are most frequently determined by 
cdaphic factors, either diree'dy or indirectly. It is obvious that greater selection 
pressures are operative at boundaries, which are usual y much sharper than 
climatic boundaries, and r.ny mutations of edaphie adaptive value have an oppor- 
tunity of avoiding absorption fn panmixy by migration, to different soil type*. Ill 
this way ihey mav escape disintegration, even though ikcy po^ser-s no sferiliiy 
barrier. Cain (1944) has summarised the earC for tht* type of differentiation 
extraordinarily well, arid concludes ''lhat speciatiou is usually a population- 
periphery phenomenon which gains ex'pTesjdon through migrat : on that allows 
i-solation and selection.'" 1 When the total peripheral areas of widespread species 
Tire considered the enormous opportunities for achievement nf ecological isolation 
nre realized. 

"There are. therefore, several ren r -on^ for believing tliat a fnirlv rap'd evolu- 
tionary devvlopmeu! of some species groups woud have occurred subsequent to 
coTitnielion during aridity. 

Some of the chief difficult ""ps ot the Australian erologist. cr plant geographer, 
arc concerned with biological diversity within ?pc:cics T and thz fact that this varies 
enormously. The range of biotypieal variation *u l rmtt(ed in a species is largely 
subject m certain taxoiiomic conventions arid the sy^iemutist's personal interpre- 
tation of them. The example of A?or'<u So-t^rnii and 4<?&fla iMtieri us a case in 
point. Here relatively minor dltT-ere^ces have led to acknowledgment of two 
separate spcries. In the author's opinion c ' () it is extremely doubtful whether the: 
dilferenrcs between these two arc beyond the bintypicai Tango of a norma! species. 
The fact that ;hc real distributions at? isolated from each other is quite under- 
standable on historical grounds- 

O The examples arc too few at firtfACUj to h-? rrncidprpd an enliYely reliable inrlcx 
uf likd> ra*ps in nature. 

(*> Tlvrn if .specific rank were jutiifleU in this case other examples aie veadih-' at 

Difficulties of this kind are common for the eealogists and arc due, amongst 
iithof thmgs, to the Tart that a italic system of nomenclature (the Lbmaean) i> 
beiu£ applied to something dynamic. For a number of practical reasons, wcD 
summarised b_y IluxJey (WOj, this is unavoidable. On the olher hand, as 
Turrill (19-10) points out, laxonomists emphasise, unintentionally, certain 
diagnostic characters wkhout lowing their real significance. Obviously, many 
problems for the taxouomist can only he solved by a consideration of the devel"p- 
nienl hf the group. Coiilitiued reference has been in preceding section- u- 
I fog \Wc level o\ competition in the early stapes of re-colonisation. This low level 
of competition lias been of considerable importance in influencing expatision of 
area and the distribution of certain species. U has been pointed out by Worthine,- 
ton ( PMO) (hat in studying eases of dirtcrentiation in nature the evolution of the 
environment must be con^idere-! in addition to that of the, and that ibe 
existence of numerous unoccupied niches and the absence of predator* arc very 
imporl-attt in determining (he amount of differentiation, possible. ,i le was consider- 
ing tltfi evolution of fresh water fish, bm hi? comments apply no less to plant 
specie.-,, and in essence merely amount to lack of coiupcli^ou increasing the ttctft 
of potential dirlcreiuir-tiou. 

Wiui the rapid expansion of the flora following flic release of climatic 
pressure, normal rates of differentiation must have been greatly exceeded for those 
bioty[)es which were suited to exploit the new edapho-clhuatic conditions, A large 
percentage of the first v*ab'e seeds of migrants would themselves become suc- 
cessful migrants, and more generations would be passed through in a limited time. 
As mentioned earlier, the brake of competition would soon he applied, but. this 
would not be fully operative-, at least for some species, until something approach- 
ing the tfSUal equilibria of plant communities was achieved, hlcre then is another 
reason io anticipate relatively rapid differentiation within some groups of the 
Australian flora. 

With relatively short-term geographical isolation of the type suggested during 
the Great Australian Arid Period, it would be most likely that differentiation only 
rarely proceeded as far as full speciation, at leasL in the sense of rhe attainment 
of inter-specific sterility-. Kxpansion of the flora from the centres of survival 
would almost certainly have led in some cases to consolidation of gene A r aria- 
tions in the attainment of full spee-;atlon. Lj :u tally, however, .such a level of 
differentiation would uol be reached, and the result would merely have been con- 
siderable hybridization. Indeed iu either case we could oped the production of 
a larae number of heterozygous hybridised and inter-crossed forms and vcty 
variable f! species complexes." Much of the differentiation developed during 
Isolation has undoubtedly subsequently been preserved by some form of ecological 
isolation. Cain fcfts [jointed ou1 that hybridization and the production of hybrid 
swarm* is particularly likely to occur with a change in ecological conditions, as 
with mares etctivities in removing a barrier and permitting free intermixing-. That, 
thi-s barrier is frequently that of biological competition is certain, but it may be 
any barrier to cros---feriiIis!lion. In the case of insect pollinated plants it might, 
for example, be du? to invasion by insects which cover a wider range. In the 
hybridisation between two spec : es of Solichufn studied by Goodwin (19.37) it was 
due hi opportunities for migration presented with the drainage of the swamp. 
Although most studies on hybridisation have been made in relatively small areas, 
it is obvious that possibilities for its development with the release of climatic 
pressure following contraction, and isolation of the greater part of the Australian 
flora were exceptional. Further, these werv greatly stimulated by the relaxation 
of another normal ecoloqac barrier in the Inv. kvel of competition, 


Reasons have been given to indicate that expansion during rccobnisation gave 
a potentially variable species, whether new or relic, a great opportunity of develop- 
ing that variability under different selective pressures, and where different oppor- 
tunities for inter-crossing occurred. This probably also encouraged an 
extraordinary biotypieal variation and the production of wil.e^prcad hybrid forms. 
Tf the. historical sequences and their consequences which have been pustulated ate 
correct, it would be expected that: (1) there would be a large relic element in 
our flora (examples have been given to substantiate rlrs); (2) considerable 
differentiation is likely to have occurred in some groups iurmg isolation, most 
of which, however, did not develop reproductive isolation; and (3) because of 
(2), and the subsequent release of climatic (and biologic) pressure, extraordinary 
opportunities for hybridisation ami intercrossing occurred during migration. II. 
as has been considered, the artdilv was centred as rtefcttfty as 4,000-6.000 vears 
ago (or even though it were considerably o'der than this), the expansion of the 
tlora is sufficiendy recent to have preserved much of this supposed complexity in 
some groups. It might be further expected, therefore, that it would be exceed- 
ingly difficult to say where some species should begin and/or end. Thai tins is 
so is apparent to any urconomisr or eenlopst who ;ub been faced wi;h the necessity 
or classify ire; related Australian species. It michr be imneipided ay a corollary 
thai: many ''species/' frequently ia:conomiai!ly good rpecics utumi no: have 
developed genetic isolation during geographic isolation and would he largely 
preserved in different areas today By ccologic b:trriers. These species would show 
free capacity for hybridisation if brought ftigftU^r. Wry lilt 1 e has been attempted 
in the s*:ud> of hybridisation between related ftpedes iti Aus.ralia, it lias been 
suspected for a Jofig time that the genus liuatfypttts had a great capacity for the 
production m polymorphism in thi? way. S"ine most in, cresting and valuable 
recent work of Brett (1-J46) has demonstrate:! Iftftl tins is so, and to a degree 
never previously imagined. His work also indicates qu;te clearly ihnt many 
species o\ Euatlypts are preserved as entities solely on account of die ecological 
isolation they enjoy. The complexity of forms in nnu:y species groups in ihe 
Australian rlora must be considered strong suppor;; ior a very rcceivl expansion 
uf area, 

J'erhaps the hes! example of biotypieal complexes recorded fjj Australia Is 
afforded by the /tumJyj'iu,* oleom P. Jncmssala p'oup of ma lock Herbert (H.'ZS) 
tcpmls (but of 53 euealypt species recorded for the gob I tic ds region or Western 
Australia (d4 endemic), n'most half the forms & are al'icu !o rbe above two 
group*. B\ (fumoxa and 325 tnvrm&ata form a somewhat Mm'tinr complex in South 
Australia.. Ann! her example of a. complex within one gfullp ot closely related 
species is shown by tbe endemics 0.1J Kangaroo tsland. Of the £2 endemics 
belonging to 4*gen&3ij listed by Wood (1930), no less than twelve are speeds find 
varieties of tbe genus PuUcucCa. Tbat is. although the e'idemic= of the region 
belong to 47 different genera, 15';£» of them are 'n one t^cnus. and most of these 
are closely rc'ated. It h apparent thai historical ecological stresses hrve per- 
mitted potential variability in the jjeiuis PhUcvluh:. on Kangaroo Island to develop 
Fairly frv-ely. As a result Pifftrjiaca is a virile genus. 

Spedcs complexes ex tl 1 1 *- type are ou'v likely In be ihoroughly understood 
as a result of cytogenetic studies, followed b> experimental analysis of hybridisa- 
tion of the type carried tun by Goodwin on St>!iclu(jo> aric being done by Urett 
tor Eitct'lypftts in Tasmania, 

( A> ifS^V ox th&xf arc obviously not deserving cf Ip^ciGf txr\.. 


4. Some Revision axd Further Considerations 

In the preceding sections a survey has ]y£$n attempted of the major geological, 
pa'eocliniatological and paleopedological factors which have inlhtenced the 
development uf ihc vegetation of South Ausiralin. Although geological eoudi- 
lions shice the Cretaceous have been fundamental in their modification of Ihc 
floral mosaic In various ways, a period of severe aridity in the middle Recent 
which followed a pluvial epoch, and was apparently rapid in its onset, has been 
the greatest modifying influence insofar as the distribution of the present units 
is concerned. It resulted hi hoth wholesale destruction of many .species and a 
contraction of area ot almost all those surviving. 

The contraction in area caused various degrees of isolation in the centres of 
survival, arid with subsequent climatic variations there has been ample opportuuuy 
for reunion; these were factors which made lot diversification. Other influences 
also probably encouraged speeiation where the potentialities existed. There arc, 
however, obviously large relic elements in the present flora— perhaps Ihc largest 
portion, and mairy of the restricted species arc old and noi new endemics. 

It is hhely thai destruction of the pi-c-arid flora frequently meant extinction 
in one refuge area and preservation in another nearby. The absence of a species 
in one region, therefore, clearly does not indicate that it never extended so fur. 
Frarueully it was merely unable io obtain a. suitable survival niche in one locality. 
but able to do 50 in another. Cases hi point are the absence of the striugvbarlcs. 
Eamfyptits ubliqnn and /*.'. Baxlcri, or the cup gums /-:. c»su>ophvUu on liyre 
Peninsula. They are present hoth on Kangaroo island and hi the Mount Lofty- 
Fleurieu Peninsula region. One would expect the southern portion of Evre 
Peninsula to be a poorer centre of survival than the other regions, because H is 
poorer in habitat diversity., and there is no valid reason lor assuming that these 
species were not ntice present there. 

Although it is always difficult, aud at 
present frequently impossible, to reach 
a satisfactory understanding of the 
reason for a particular species distrihu- 
i;on, fc-c c?n be relatively certain of one 
hnporOmt fact. Thai is that the plant 
t oatwuntties themselves are \ouug — 
the combinations ate new. One would 
vx pi rl as a result that they would 
show some signs of instability. There, however, no great in liabilities 
apparent, and this must be due to 
j »e enormous c iahiJising influence 
ot p'.K.'-C'-sioii, together with tho high 
coioni-mii* potential when the level 
of competition is low. The first 
oceu;:ai:t of a vacant niche therc- 
ai'tei holds a great' advantage. One 

case in which the relationship beiwcen two associations appears to be in 
a state of ih)K \* that of the Jiiu'Qlyptnx dit'crxifaliu association and the 
Melaleuca - piibcscnts-Casuarmu sirirta association, where the two occur on 
shallow terra rossa soils: developed on aeolianite. limestone- Indications are 
that the balance between these two associations, the former a sclerophyh scrub 
and the other an open savannah woodland, is a very delicate one* and that the 
sclerophvll is invading the woodland in smite places. At Section 6, Hundred Uley, 
Eyre Peninsula, the area occupied by the sclerophyll scrub today is very much 

Hl'fOnCO OF *V***iLUA 


£& \2 

Enculyptiis (Jk'crsif^lta association (fte(9ttc4)i 

Section <\ Hundred Uley, Kj re LVniusula. 

The f.h'jnblc-hat^lied portion is fln exttusion 

since the original survey. 

grpaki Htan that rlclimited by the surveyors in their land survey nj IiS77. A 
cft&CTJCnt type of instability, which is ppiehlisd rather ihao real, is shown by 
specie^ which are growing at tire extreme limits of their lolernnce-;. Must of these 
must he poor competitors and could readily be replaced bv apfifcifiS more fcitltgd to 
the environment were ih*ry available- Fxump.'e.- of l|pfi n; fnre include; Lucalyl'ftts 
cltuiocolyx on EyW: I'e.ninsula, which is existing over much o[ its area in a 
depauperate state on Bttfljfe and under climatic conditions which are supporting the 
more highly integrated ZL Baxlcri a^ociatlon on Kangaroo Island. Another ease 
is the extreme dwarfing of IL Bn:th:ri ai fhe limits oi its cdapho-cUmalic range 
in the Upper South-East, and then there are a lar & r e mtrubcr of r-pecies which occur 
in an excessively dwarfed state in many 6f our comimmitnts; for example 
Batiksio warffiuti'ta, which occurs normally us a tree or In'l .-hruh, can be reduced 
to less than six inches in height and still eceeise. 

Bern use of the complex of factors which can modify the relationships between 
soil and ve^eintion it is not surprising that other things are not equal, and the 
generalisation that wiLhtti Lhe same climatic horizon the dislribulion of vegetation 
is dciermined by soil conditions frapicutly does not hold. There are difficulties in 
defining- soil-vegetal inn relationships precisely. Firstly, there are the laxonomic 
problems mentioned earlier, and secondly there are, from :hc Sail viewpoint, the 
severe limitations imposed by analytical ter.hnkjucs. especially those which attempt 
to evaluate soil fertility. These difficulties will always be apparent because &oil 
fertility, for instance, is without exact definition unless referred to a particular 
blotvpe or series of hiohpes, wbile the subtleties of geuotypical variation which 
have ecological significance will always escape lhe taxtnomist to a greater or 
lesser degree. Added to these problems are fhe difficulties of assessing the 
climatic and purely biological factors of Lht! environment. Indeed, plant ecology, 
or for that matter CcoiogV generally, is faced with some very real problems.. The 
acceptance of the fact ihat specifitJOT represents no absolute stage in evolu- 
tion, (see Mailer 1940,1. but is graduaijy arrived at. arid that in the Uixonomists' 
species all types of intergrades occur, means that identification, and understanding 
of the gmorypieal variation with which the geographer or eco!ogi*t is dealing is 
a necessary prerequisite. Further, he has to assess tbc complex environment and 
inter-relate these two, that is organism and environment. Although in a general 
way the factors concerned are known fairly well, further progress so far as plant 
ecology goes must depend largely upon progress in other fields, such as pedology 
and climatology. 

Despite lhe limitations imposed hy difficulties like tbc above, and the dis- 
continuities Ibat have hecn stressed, it is: nevertheless possible to arrive at very 
useful ctassifiriil inns and understandings of the vegt-tai ion. South Australia has 
been a good centre for working ont many of these relationships because the Overall 
simplicity of the soil mosaic, the regularity of the climatic zonation, and rhe limited 
X&hlgtSbs both hi extent and number, from which the present communities have 
developed, has impressed upon the vegetation patterns a sirnpieity that is wanting 
in the en stern Stales. 


The dynamic view of vegetation has resulted in the concept of a succession 
of plant communities towards a rl'tn^Jx association determined entirely by climate- 
Of the earlier propoundcrs of the succession concept perhaps Co\v!es (1901 ) was 
lhe fus... Ft was Clcmenls (191o), however, who brought it prominently to the 
fore and e'aboratcd it iu his publications until plant eco'ogy '.^camc burdened 


with a very special term i no logy, chiefly developed about deductive theories. The 
succession of associations leading to the final climax is known as a serf. It was 
suggested that in the present-day communities the climax Vmfi readily recognis- 
able—it was the most widespread assemblage witinu a climatic zone ami that all 
other communities could be classified as sera! stages, either as relic corrnnuniric? 
or as 'Communities leading directly to the climax, 

A pioneering attempt by Wood ( 1937> to classify the communities of South 
Australia and show their mter-relanonships on a successiona! basis, although dome; 
much :n brings order out of chaos, wa.s soon found warning, it resul'.ed in a 
reconsideration (Wood 1939) of some of the fundamentals un deriving ecological 

Tl'H failure to apply successfully the succcssioHul concept to a clarification 
nf Soii.ii Australian communities has largely been duo to the hi*-torical develop- 
ment of the vegetation. We have suggested that the presen: associations have 
result e I from the re-colouration pf vast hare areas by dispersal from centred 
of -.111 viva*. The associations are very young and their distribution has been dete- 
mhio.d uiLhin a climatic zone ch:eJly by edaphic conditions, i Ji<- distribution 
patterns have been greatly udhumexxh however, by other factors such as individual 
dispersal capacities, chance dispersals, oppnsin<r migfEUiQU& location of survival 
centres, biotypical dillerentialion. barriers, ele. it js ntft surprising tft&t Ihe dual 
spenes-aggregates ( associations, etc.) are not nude up of tmbs with identical 
tolerances, nor would this ever be expected. Although, therefore, the species 
which p'ayed a dominant part ir\ the developmental seres a>soc:utel with re- 
•coloniscuiun are probably all still living, it is obvious that present-day ae^regaie-* 
are different and cannot be considered serai stages in the accepted Clements sense 
— un.ess. indeed, thev all he considered edaphic snbclhnaxes. One might as well 
consider them all edapho-ehmaiic climaxes, but nm'-t appreciate their dvuamic 

U is a logical consequence of speciation an*! evolution, and of historical 
changes, 'hat the dynamic nature of vegetation wi«! always he nraiiitained The 
decree of stability will be greatly governed by the historical sequences, and these 
will vary from country tn country. It i^ perhaps understandable lhat the concept 
of succession towards a crmax and the concept of ebmattc-indueed migrations 
of whole communities, rather than individuals, has been pushed so far in America, 
-where l/.e paJcoutulogical record indicrifes a very fang history, and when 1 climatic 
fluctuations, though extensive, have been gradual, Cde-ascui ( 1 r >23 ) , for example 
has been able to produce a con.dderabk' tpuiruity fof data to .show that the principal 
vcgcLatienal elements in ti;e Middle \\r : ,\ were differentiated in the Tertiary and 
"have contmuousy maintained rheir pre-cut relative position. v ' Such commuily 
of velaiioie-hip could never he envisaged tur elements of the Australian vegeta- 
don. in the Tertiary, when Australia was redure-d to an almost perfect peneplain, 
eHma'Ic climaxes, would perhaps have been recognisable in the Theoretical sens*. 
The ecological concepts and units of classification proposed by ClcuieuN 
(l r »16d, and followed by Tandey. ^ere based upm Ihe philosoxbaxif eeria -^ of 
organism. Since this ciasdhcation had been found unworkable for South Aus- 
tralian vegetal ion, Wood ( 19.W) propped a rational basis which should underlie, 
atn system td* classification n|' plant communities, and tentatively ^iggested units 
of convenience. It is proposed here to amplify and expend the cousidcratioits 
given in the previous paper, and, -n the light of further field experience, to define 
units suitable for classifying vegetation. 

Ecology, in large measure, consists in denning the hunts of species which 
grow nuturaly together, and in understanding the factors responsible an* their 
maintenance ae a community. Ecology, afier all. Is a branch of physiology.. In 


the laboratory we study the reactions of species when all environmental factors 
save one or two are constant; in ecology in the held we suidy these reactions when 
the whole constellation of environmental factors varies and where we have the 
added complication of competition between species. 

The basic tact Underlying all ecological work is ihe mailer of experience 
that any particular species will develop and be maintained only within certain 
environmental limits — these limits include soil conditions of nutrients, water, etc.. 
and rlimatie conditions. These limits we may term the ''potential environment" 
of the particular species. 

Any particular environment selects from any population of species exposed 
to it those species whose potential environment coincides in part with the actual 
environment — in other words, we are led Lo a simple Darwinian explanalion. 

Competition between species and changes in the genetic constitution may 
cause changes in frequency, and an initial assemblage of species may ahe*- the 
v chemical environment, so opening the 

way for further change in the speeir.s- 

The potential environments of com- 
paratively few species arc accurately 
known. Wc are familiar enough with 
species with exacting environmental 
conditions in tie case of bacteria — the 
basis of the technique of bacteriology 
is lbs prodverou of selective media 
for growth. Wood and I'aas Becking 
( i^3?) have drawn attention to fcfec 
miivfTf-.a! occurrence of a community 
of gri5£?jr.j bhie greens and diatoms and 
Amu mated bv Kttbpm niaritima in salt 
hki-$ where :he concentration of salts 
is 6-lC c /o. And in a more extreme 
environment, viz., salt brine* (up to 
$my\ 2Q c /r. iota 1 - salts), there appears a 
coiuuumity identical in all parts of the 
world dominated by Dvnaficlla rhidls. The potential environments of members 
of I his community for three, chlorides al different rorceotratJons have been 
delimited by Kaas Becking. 

Wood (1937) has drawn aUeu'.ion to convergence in species-composition of 
ohgofrophie hogs (pH 4*0-4*5). at D rem he ( Holland ) and Mount Compass 
(South Australia^ v/hcre c'oscly allied species d£ the same genera occur in the 
same communities. Eardlcy (1943) studied a Somh Australian fen (neu'rul to 
alkaline prats) and showed ihe remarkable similarity in sp-ecies-conrpasiLion with 
Kast Anglian fens and some North American -wamos. Wood (1939:) illustrated 
how* some of the chief associations in South Australia were related lo annnai rain- 
fall, 2>hosphate content and pH of the soil. 

It :s a matter of experience that some species have a wide potential environ- 
ment, whilst others have an extremely limited one, and tais is particularly true 
of many species of the genus Eucalyptus. With these underlying ideas in mind, 
vegetation units (/.c, species-aggregates which live naturally together) can be 

Suppose we consider a hypothetical case of a community of four species, 
a, fat i\ d 3 whose potential environments with respect to two independent factors. 
say, .r and v. are as follows, the species a extending over the whole area (h*g. 13 J. 


y \/ \ ,--\ /, x a 

fife: 1.1 


If the unit community is defined as one of definite and uniform composition, 
as was done by the Third International FSorauieal Congress, then it is clear that 
there are several different units occurring within the habitat. These units are 
the "associations' of Swedish Geologists. They lead to a completely unwieldy 
classification, r.^ v Ostvald (1923) has described 164 separate associations in a 
stretch of moorland 40 square miles in area. 

On the other hand. Tansley (1939) considers the tree dominant only as 
denning the association, and on this classification the whole area dominated by a 
would he the unit. Tansley f s classifier:! Ion of the British oak-beeclnvoods, together 
with some habitat factors is as follows: — 

Association (climax) ; oalc-beeehwooi 

Consociations (one dominant only). 

I Qitcrcclum rohoris - - damp clays and loams: 

oa k 3 neutral to alkaline 


Qunxcinui sexsilifiontc - (a) savannah wood^auds-podsols 

- (b) with heath-peaty podsols 

SFagctum cakicohtm - - rendzina pH7~5-8'0 

Fngetum- nibosum - - brown earth 4*5 -7-7 

Fcgpfnm crkctoswn - - podsol 3*5 -4' 5 

concedes (serai stages) are; 

birch wood — — > oakwood 

ashwood — — > beech wood 

Tins unit of Tansley (the association) is too broad a unit of classification 
for practical purposes. Indeed, the example above seems to violate the very idea 
of an "'association" of plants, for the groupings of associated plants arc totally 
unlike in the different communities — some consociations have no species in 
common with each other! It also violates Tan sieves definition ''constant habitat," 
for, although climatic factors remain relatively constant in the above association, 
other equally important factor?, wfe.j soil types, nutrients, [ill and water relation- 
ships are widely different. The system really breaks down owing to the wide 
potential environment of the dominant species — much wider than that of any trf 
l lie species associated with it. 

Suppose we take a specific case, and apply Tanstcy's classification to the. 
communis ies in South Australia dominated by Eucaiy^lus Baxtcr'i, the brown 
strlngybark. The following well-defined communkies are readily distinguished: 

E. Bv>xteri-E. Hubcriarm - - wet podsols 

E- Baxtcri-E. obliqna - - - normal podsols 

E. Baxtcri - dry and shallow podsols 

K. Baxfcri-E, divcrsifolia - - siliceous sands and 

residual podsols over ironslone 
E. Baxtcri-E. cosmophylla - - latentic soils (residua! podsols ) 

The rainfall over the whole group of communities varies from 40 to 14 inches 
per annum. The florisdc make-up and species-frequency of each of the above 
coumi'-nhies is quhc* distinctive; some of the communities have few species in 
common except E. Baxtcrt. The first community is a savannah woodland, the 
second and third dry sclerophyli forests, the fourth a mallee scrub, and the fifth 
a treeless uuiechia. There is no evidence whatever of any successional trends 
between them. 

Furthermore, in the South- Rasl. of South Australia, ranges of residual 
podfnls occur parallel to (he coast, -and at ri^ht angles to the rainfall isohyers f 
from 30 inches in the south to 14 indies in the north, Proposing northwards 
ofic finch a gradual dropping oui: of seme species and their replacement by o l hers. 
/:\ Boxicn continues as a dominant throughout ; in the welter area* as a forest 
tree, then a* a stunted small tree, then as a shrub less than 3 feet high. Ftnally 
it disappears, hut most oS the plains associated with it in the stunted pha^e con- 
tinue on as heath or mallee-heath. obviously related to the former community. 

It h clear that h. tlnxicri is a species with a wide potential environment, 
much wider than that of most of the -other plants associated Willi it. To delimit 
a community bused on the distribution of R\. Haxfcri a!onc is to ignore the asso- 
ciated species and simply to define the area occupied hy the cuealypt species. 

Relatively few species are distributed purely at random, and associations. 
under whatever system, arc determined subjeaive'-y. in defining an association 
we i?Mc a middle course between the British-American and the Scandinavian- 
Swi-'s schools. We define an association as a constant associat'on (i.e., growing 
totjethcr) of dominant sfccic.s recurring in similar Imnljis. The term dominant 
species refers not only to the tree species, but also to the dominant shruh and/or 
herb spgripa which give the characteristic look or facie- to the community, It is 
similar in practice to die "consociation" of Tans'.ey, hut without the implication*, 
inherent in his definition, This is the gfuttpiiig of greatest (tie in vegetation 
studies in Australia, though not ner.cs>arily the most 1 urdameutal unit, hi prac- 
tice we have found associations to he closely correlated with soil types; Of if on 
different soil types, then some compensating factor can normally be found, e.g., 
water relations, nutrients, etc. 

The association is made up of smaller units and may be grouped into larger 

The smaller units arc the type and the society, lypa we define as local 
clunu;r in the dominants of the vppcr siraimn of an association Tt'htcli is accom- 
panied by little or no chumjc in the other domhnuils. A Society is a local change- 
-\n iUc dominants of the loiter stratum. 

"he concept of type has been used for a long time in forest praelice. Appar- 
ently it was first used 'by Graves (1899), who staled u lhc same lype of foresL will 
tend to be produced on the 5-ame classes: or eiiuuuiou and soil in a specified region. 
There will be variations within Lhe type, but the .:hanicleri?cic features will 
remain romtant, that is the predominant species, density, habit ot trees, character 
of undergrowth* etc. ti aponion of the forest is destroyed by fire, wind or other- 
wise, the type may for the time being be changed, but if left undtsnnbed will 
rcver tQ the original form, provided the coudition of Lho soil is not changed. SJ 

This definition, in essence, is sail retained in the Glossary of technical term* 
of the Society of American Foresters (1914), where h is added: "The icrnt 
suggests repetitions oF the same, eharaclcns under similar conditions." 

iTyor (19.59) first used the term in ecological liierature in Australia, usin. Y 
"quantitative floristic uniformity of the dominants" as the criterion. Pidgeon 
(1942) defined forest type as "a forest stand which has. wherever it occurs, the 
same fioribtic composition of dominants, and which 'develops in essentially similar 
habitats." Tn both definitions the term "dominants" refers to the dominant 
tree tpecie.s. 

1'otli tht above workers have used tins unit with advantage, for it is a reality 
in the field. The authors of the present paper disagree with their definition of 
tores' type since it neglects the associated species, and in this way departs some- 
what even from the original concept. However, the difference is one of definition 
rather than of fact, for Pryor live, eit.) states: "Jn an area with very few 


dominant species e.g., jarrah region of Weston Australia, the lowest order com- 
munity {ie- f type) must be determined by some criterion additional to the floristic 
composition of dominants, c.u lt the (loristic composition of the lower strata of 
the community" and again, "the vegetation type corresponds approximately with 
the Association of Kraun-Blanquet and the Faciation of Clements." The essence 
of Braun-BIanquet's association is "uniform fluristic composition" — not of 
dominants only. We prefer, therefore, the definition ot type as described above. 
In practice, there is no conflict between different Australian Avorkers in actual 
delimitation of types in the field. The concept of type has been used in studying 
the forests of the mountainous areas ot New South Wales and South Australia 
(Pryor \9W\ Pidgeon 1942; Hoom-rma 1946). in these areas the need for such 
a unit arises from tbc extreme sens ; rivity of many Hm-atypias spec>s to chants 
in the micro-habiUU ; ctumgfs which in oilier parts of the worUl cause alterations 
in the shrub or herb layers only. It £g clem* thnl some eucalypt species have ;i 
mote reslricted potential environment than have the *hrub species with which LliCJ 
are associated. 

The type is a more fundamental unit than the association as dchued above. 
and it will be clear that an association as defined by ikt as a collection of types 
In a forest association, the floristies of tlie associated species of the under; -rowf.*. 
rvmabi the same, though there may be local changes in the tree .-pedes. 

In urnling associations into larger groups ftiy characteristic oi the association 
may serve as a basis for classification, These classification*; abstract same 
charaeterisftcs of the association. Ft is clear that several alternative classification r. 
can be made; they are not necessarily mutually exclusive bui are complementary. 
and any may shed some iight on vegetation problems, 

The French-Swiss school gfQUJW us-'Ocinf-ons into larger units on the bask of 
a common Mora, although (his sometimes leads to £roupin«: together of association- 
whtch are not nearly allied. Using structure and life form as criteria, association^ 
may be grouped into .such units as seJcrophyll forest, savannah woodland, savan- 
nah, etc*, in any particular area. These are the Formations of Tunsley fcifd QJC 
grouoin'j.s of grca? convenience. although &eir use does not always lead, (jo 
grouping of nearly related associations, cji., the mallce-hcath and heath of ili»- 
upper South-East (Jessup 194n) r nearly related in fieri s tic* and habitat would he- 
separated on this basis. 

In South Australia we have used one feature of the habitat, vi:<. t itllied soih:, 
ossr'Cfafrd with yrlaicd ftori-siic composition of spveios as a basis for classiuciUinu. 
Whhin definite climatic limits, associations on nearly related soil types and with 
related florist ie composition are grouped Together as an F.i/aphu: Cnwplcx. \ 
striking" example ot an Edaphic Complex Is the associations found on pod^ohV.ed 
soils in the -Mount 1 ofty Ranges within the climatic zone limited by the 25-50"' 
annual isoyet. Detailed work in progress ( Spevhr VM6) shows that on tin- 
soils weh-difined associations occur with the following cucalypts as dominant; 
/:. nbliqiMr /.'. Buxi-rri, R. CdSfnopltyila, IS. fasciat!of>>, /•'. l t 'tu:oryh>;>. The 
potlsols include normal, immature and residual pounds, podsuU on deep >ands 
unA ,( t;re>-l»i'o\\ n" podsols ; they vary iu tlieir tiulrieut and water relation?. 
Floristic lists tor each of these communities show xkat wilh lew exceptions ilu? 
same specie:- occur in each of them; llieir frequencies^ however, differ greatly in 
the different associations, hut arc constant for an v one community associa.ed with 
a particular eucalypt. Shrub species dominant in one association may lie only 
occasional or rare iu another. If is possible that here we see the selective effcci 
of environment at work on a relatively yOTJtng species-population. Selection foft5 
proceeded lo such an extent that the separate associations can readily be dis 
tin^an'-shed, hut tbc associations themselves have unity in floristic composition and 
in habitat m the Kdaphie Complex. 


The greater portions of South Auslralia possess a heritage from the past, an 
thai manv soils retain certain morphological features which are "fossil" if., not 
Eormecl in equilibrium with present climate, but rdieS of former, more or less 
irreversible, soil processes. The result of Recent aridity has been a stripping and 
resorting of [be upper horizons of many of ihe old soils, and the building up and 
exposure <& ^ftorf* ^ material, which tp now rc-subjected to pedogenic processes 
The result is a very diverse pedogenie hislory for many .-oils. 

In this complicated variety of habitat? tlte ftdaphie Complex is of value. For 
example, in the tipper South-Eash on neutral to alkaline sands, Jessup (1946) 
has shrrwn that a malice scrub — the E. divcrsifolw-L. attgittosu association — occurs 
on crcsly of hills; on shallow sands pT the same upe, and on well-drained adjacent 
:-and plain or heath, the Ctniiitrina pusilUi-Xttntttorrhocv nusiralis association 
Ofcdl '';•'. The hill slopes ft an cartons or transition region wiih dominants from 
both r.^ociationi? present and with F, Irplophylla more promineui- ■ ibis is the well 
known malice-heath. In the two agnations the associated species are quite dis- 
tinct. 0ii undulating plains in this urea the vegetation r.t Sist sight is extremeK 
coniph'N, varying with every rise\ rind its edaphic relationships only became clear 
when the well-defined sandhill-plain region was studied. Sue!" areas can often 
best be mapped as a Complex and sorting out of associations left to detailed study. 

Similarly, on the latcrite and associated -oils on Kangaroo Isiaud, the fol- 
lowing associations can he distinguished: 

11. Baxtcri-E. cosinopkylla - - on lateritc soils 

/J, remain - ... ou heavy or massive lateritc 

li diver sifoliu-E. cosmophyllxi - on latcrite soils with some 

siliceous sands 
. r BaxUri-E. diver slfolia - - on siliceous sands 

(A horizon of the laterite) 

The ma!lce-b**ooir;bnsh complex in the Sonrh-Kast. Kangaroo Island and Eyre 
Peninsula (Crocker l<H6, Jessup 1946, Wood 1937 j is a good example of a 
eomp-c* of associations closely allied, but showing soma marked floristic difTer- 
euces owing to cndcmUm on Kangaroo isiaud and Eyre Peninsula, and to differing 
selection pressures in centres of survival in the three regions during the aridity. 

The vegetation on the immature soils derived front the Bawkesbury Band- 
stone in New South Wales, and described by Pidgeon (1942J, is an Kdaphic 

In the associations described above there is vo evidence of succession or of 
invasion, but only of transition. Indeed, selectivity ot' environment — especially of 
climatic factors — is particularly marked in sev<ra1 areas of South Australia where 
a relatively uniform soil extends over a wide range of present-day climatic condi- 
tions. Examples are the ranges in the South-F.ast. the residual potlsols on Kan- 
garoo Island and Ihe. malice soil?. Here we nuiy follow the slow dropping out and 
replacement of species by others as rainfall decrease?, tilt at the two extremes are 
found communities with few or no species in common. 

Commimitics which in Tansky's system are regarded .as- serai, i.e., those which 
show allogenic succession, nuty also be grouped as an edaphic complex. The 
communities on steep hillsides and skeletal soils, as well as on mature soils on 
itOpS of ridges, and on gentler slopes in the Mount To'iy Ranges, may he cited 
as au. example. In each case the habitats are different and selective, so arc the 
floristics, although they arc allied. The hilhiidc communities are stable, and a 
unifcrm terminal community is only ihcorctically possible b_y a levelling down ot 
all physiographic units to the uniform soil type. These examples might be termed 
edaphic successions: but in any case it is desirable to separate I hem from biolically 
induced successions. 


The examples given of the past history of South Australian soils and vegeta- 
tion and the in^punsiJiility of reconstituting l!ic past, illustrate the dangers of 
dogmatising- about successions of this kind. All our experience indicates that 
succession should not he used in any scheme of classilicaiion of communities. 

This does not imply that biotic succession does not occur. It is ohvious on 
sand dunes, and in swamps generally where the plants themselves cause changes 
in thv:- habitat to produce a special soil carrying, at equilibrium, a terminal com- 
munity. Within associations as defined hy us biotic successioual changes occur 
following tire, felling, grazing. etc., there is no evidence rii serai changes, 
only a selectivity of the habitat, This is well illustrated in die lower South- East, 
where the fallowing conrmunines in the Fame climatic zone are clear-cut both in 
Romtfcs and habitat and wi;h no evidence ol scral stages between or leadira r 
to them. 

/:. ll'xferi - Sclcrophyll forest - - on re? i dual podsols 

Xanilmrrhoca-Hakea Heath - wet acid podsols 

Ji t Lanalilulet^is - Savatmah woodland - - meadow podsols 

H. ovata - Savannah woodland - - intermediate meadow 


if fili 'tvt-Cladium - Savannah - rendzina 

Ahti'Jcuca pubfscens Savannah woodland - - terra rossa 

Much ox the lack of succession is doubtless due to the low degree of integra- 
tion ul the open and licJtt-dL-mandm^ communities of Eucalyptus as well as to 
the historical factors discussed previously. Invasion is more obvious in highly 
integrated ci'inntuuities atid eerrainly occurs in Australia where rain forest 
iurpinge-. on Kucalypt forest, but the lack of succession in our forests generally 
prohibits i£s use as a characteristic in any -cheine of chi c chicaiion. 

Of rhe dements which make up the Australian flora the Australian element 
is the most prominent m the southern regions. As has been, indicated its centre 
of origin tq oh? c tue. hud in aU likvlihoocf there were many centres. Its present 
distribution Ktsfrtyftg rai^csN a southern origin, but it is mure, likely that the 
primary Yemenis fov^td when AtMraha generally had a temperate and move 
uniform elm-are. Hcrhtrt (iv'2S) has sir^estcM ihat the genus %ktdy$iat 
successfully established itseii under a icrripcrmo climate with abundant rainfah. 
Il is perhaps s^mheant that a!ong the invasion routes of the Indo-AIelane^'an 
element^ there hay been practically no two-way tramc. This Jack of reciprocity 
\\as early tinted by Hooker f I860), and it does suggest invasion- following with- 
drawal o» the Australian eh.ment to the soiuh. hi any case, the evidence Is very 
sirou^y in favour of scmuldn- in the nature of a pan-Australian flora in ;hc 
early TerJuaJJ — at least over ihe southern hall of Australia. Such a possibility 
isniade more Hkeiyby the edaphfc and climatic uniiormhy which existed, for at 
thfa time tins continent was reduced 10 an rfmiftl perkct peneplain. 7 This 
urmomrry was broken bv marine transitions, volcanic aclivirv and carih- 
movem-nts whxh reached ihefr maximum hi the Ja':e Tertiary, thou'di each, and 
especially fhe latter, enn-fed on into ijj- Oua'vruarv to sonic exreni. Habitat 
diversity was further increased m the, FlcisK-cene bv the gcr.exady hHi rainfall 
causing- active erosion of the newly- formed divides,' and the spreading of laro-e 
alluvia] deposits, in the late Plcislocene orher factors such as the deposition of 
large quantities of calcareous loess oyer Southern Australia led to a modification 


of the edaphic conditions, while \1\ the Quah"rii;iry ^cne;-aljv, iSfgJI Oscillations 
in dtti flora occurred as a response to chafing climatic, conditions. 

Of these climatic changes,, the QMSl ^gjtificatfi from the viewpoint oi the ijis- 
Lribitiion ot the present vegetation comtnmiiics has been a pnst-I-'Iristoccnc period 
of aridity, v/hich fodowod pluvial andiiincs. 

The desiccation was so severe ;md sudden that it T ( -ultcd UTS considerable 
portion of the prc-arid ilora being- entirely aut. Tfefi surviving remnants 
were Isolated in numerous refuses. nu>M ijtj^nnawt ot w! ich U'i-;* those recalls 
where habkat diversity,, especially c! inn lie divci-ity, was ^realctf. The prcsent- 
dav plairt eomnmu hies are the resist ol re^oluiii ration it v;,sl virtually bare, 
areas, especially in the arid regions. Ifi& ttKOMffcto* achieved iLa gmfifet stimulus 
with a slight release of the climatic pn-;si.^, The Arvtrahan flw» today con- 
tains a very large pre-arKI t die element D&pltt the Efc^QVbfe enounce; lor 
biotypica! rjitteremiatioii with reunien following isolatic h and the posMbi!ities ot 
later invasions (greatly floured by :he low lev:.] pi competition), it is likely that 
tills relic portftflta still the fftrgt*. li; a broad way the JkK of the arid regions is H 
blending of the Inrlo-lVfelanoum element from the noM'i and north-east with the 
Australian element from the south-wed., south and suuoi-east, wh-'eh has occurred 
subservient to the Great Ati&trftljten Ard l';-ri.,»l |l can be looked on, ihcretore,, 
as a lar^e ecotonal region. Tins hl&i<$ug oi' the two elements is well illustrated 
by the ffnristic analyses of Murray (WA1) and Carduer {1042). 

There is evidence for a post-Pleislocene x.-roihertuk: period in many places 
in the woHfl (see Huntington and VfjSbtfr 1922£\. !ml nowhere does it apotar to 
have b^en so severe, or lo have had soch far-reaching consequences a-- in Aus- 
tralia, unless perhaps in I he southern Mediterranean region. In view of the close 
relationship between the occurrence of nuixjminm aridity, wanner seas and higher 
seas ( KJ-20 feet) in Ausiraha, and the evidence tor a postglacial worldwide 
(oustaHcA rise in sea kve.l of this o-der, it is not. unlikely that anility was much 
more general at this time ihan has been imagined. J r ans<:n (l'TU), from s.udics 
on the" Pear River delta. Britidi Columbia, hn? concluded that present tempca- 
tures have not persisted for more ihan 4.000-5,000 \ :*a* s, the mid post glacial 
filing warmer. Although Anslnilwi largely lies in the Irlitudes where the high 
pressures len.l to tavour aridity, it is likely that the widespread eifects ci the 
desiccation Ave.rc due to a combinnliou of factors. Of thcr.e the general lack of 
relief {near-pencp'anation) oi the greater part of ihc continent, large areas of 
laterised soi's which in many ami* probably preserved up to thrs time, much of 
their easily -s1 ripped A horizons, the occurrence over much of Australia of great 
quantities of ailuvmm and (lefrilal imperial, and of certain lime-rich soils in 
southern Australia which were liable 10 easy loss of stability, together with the 
presence of a very messe flora, are most import anf and ire sufficient to e:;p:ain 
the results. 

One fact which is appaicm from (he: ana\<ds of the development, of the South 
Australian plant communities b- thai, the ast-ocuiious themselves are very young. 
Their distribution has been determined wirbin a clim;tx zone chiefly by c.daphie 
condnioijs, in thai ihe.»e have been responsible for the sequence of related habitats, 
which have made up the migratory routes and utve largely determined the speed 
and direction of migration. 

A* alreadv metitioned the distrbiuion patterns have been gieatly influx-need, 
however, by other factors such a.s individual dispersal capacities, chance dispersals, 
opposing migrations, location of survival eeutics, varying degrees of biotypical 
differentiation (including speciation), barriers, etc. It is not surprising that the. 
fmaJ species-ae;g rebates (associations, etc.) arc r.ot made up of units with identical 
loleranccs. Nor is it surprising that an attempi to inter-relate these communities 


(Wood J937) as serai stages in the accquccl sduse of Clements (QcSMUR MH<i) 
should have been unpurcessiuh Tiif dfcg^CC of stability of vegetation will he 
governed \;,y the historical sequences, and these will vary greatly from country tg 
country ami within a country. 

In some respects the response ot the flora to aridity in Australia was vn*\ 
similar lo mjhu? of the flora rlucuui'iono of the Pieislocene iec-ascs in ihr 
northern hemisphere. These, ])H our arftSjy, wipcu out (he vegetation over large 
areas,, I>iit ieft centres of survival G'adation. however, was a much uiurc i'mdual 
phenomenon, permitting a considerable amount oT adaptive differentiation 10 
proceed, More important still was the fact lhai most species bad a sufficiently 
Avide amplitude and a fa%t enough migration eapaeuy to keep ahead of tlic icq in 
its advance, and to follow it fairly closely in retteah In this wav wholesale 
destruction was avoided an 1 the pattern di rc-u>]i million has been, in fact. 
entirely different. Chances for tmlimited h)l»riilisfitfijn and/or consolidation - -I 
hjotypical differentiation did not occur to the same degree. The time factor, too, 
l.as_be;n ki very drttere-nl. The historical stateness and their consequences &&'.ai 
suflk>'.:r to exp-ain the yped lie tfn&ttity of the rouli liuropcan sud American 
flocps hj comparison with ttiC JjpWfi jjuvific complexity of the Australian .'lorn. 

The ordinal and specific peculiarity Of the flora of south-west Australia and 
the high percentage of endemics (70-80%) is considered to be adequately 
explained by a consideration of the palcobounicul evidence in relation to geological 
arid climatic history. A pan-Australian do: a probably existed in the early 
Tertiary. A southerly climatic-induced contraction in the Miocene coincided with 
the inundation of lar^e portions of southern Australia, and effectively isolated 
doristicuhy south-east and voudvwest Australia. The sea had retreated almost 
lo its present position by llie end of the Pliocene Despite certain expansions of 
the flora since that time, a large measure of, if not complete, isolation of the flora 
of the south-west lias subset] uvmly been maintained. The barriers to migration 
have been chiefly cdaphic ruul climatic, though the foundering of the gulfs of 
South Australia was also significant. 

The richness in endemics of the flora of south-west Australia is the most 
interesting feature geographically in the flora of the continent. 

lr was mentioned in discussing this matter earlier that the flora isolated in 
(M • truth-west by the laie Torpury sens wa« a peneplain flora, and a hat before or 
ftityft Phtti it must have become "adapted" to laierkie soils because they were at 
-some stage of the Tertiary a widespread feature of the peneplains. * Western 
Australia largely avoided the kite Tertiary-Quaternary uplifts which refcjltsd 
in the eastern divides and the Mount Lofty- Flinders system, and has been both 
prior and subsequent to this, one of the most stable parts of the Commonwealth. 
The Identic soils are preserved there to a befrer decree than anywhere else in 
Australia. This emphasises die importance of eduphie harriers in maintaining 
isolation after the removal of other physical barriers on the retreat of the Tn- 
h'ary seas En addition to the edaphic barriers: mentioned earlier is that caused 
hv the development, with accession of calcareous loess durinrr the lale Pleistocene, 
of ft laia;e area of pedocalclc soils lying- right across the migratory routes. This 
alone wituld prove a severe and extensive obstacle to late nugraiioiis. That .the 
\uFtralian element in the south-west of Western Australians one selected hv 
lateritic soil can hardly be doubted. According to Gardner (he. ril.) those part's 
til the smith-west richest in the purely Australian, element are the sand heaths. 
'•Thefloristie weal lb of these heaths is probably unparalleled anywhere else Oti 
earth, ,T he says. I* is significant that these are in the region of Australia vherc- 
parla of the old laterilic profiles are preserved, but more significant, these sand 
heaths are ?orae of the few regions where the old A bomon;-; of (he latcrin* 

truncated more or I«$, and perhaps feltfficriijff some local rearrangement, arc still 
preserved. f,J) The sand, which is underlain by laterite, is portion of this old A 
horizon. The floristic stability of the flora of the south wfcst is paralleled, there- 
lore, by a great measure of edaphie -lability. Because of the narrow climatic 
zonation, climatic stability, or at least the provision ot a wide range of climatic 
habitats within a. short distance. Las been maintained also for a long time. 

From the survey that has been made in Ibis paper of geological history, and 
its consequences on the Australian, and especially the South Australian flora, it 
is apparent that these geological factors have played an important part in the 
development of many groups of plants. 

The iorcgoing considerations both on tbe Australian flora and the develop- 
ment cf the vegetation of South Australia, are based on only a very imperfect 
understanding of Tertiary and Quaternary geological and climatic history, in 
addition to a very restricted pnleohotanical literature. Their tentative nature 
must, therefore, he stressed. However, although many modifications to the con- 
clusions arc to be expected, the present agreement amongst geologists on many 
of the basic farts in the Australian post-Mesozoic record, extended ecological 
and penological knowledge, and the establishment of many new principles in plant 
geography are sufficient justification for tin ni tempt of this type. Further 
improved interpretations will depend largely nn advance? in Australian geology, 
pedology, pnleoholany (including paleoecnlogy ). cytogenetics and phylogeny. 

This analysis of the time facror in ecology with its insistence on The selective 
effect of edaphie factors and migrating* plant populations has strengthened the 
view put forward by one of us (J. G. W. 19.39) and amplified here, vti., that 
the basis underlying ecology te n physiological one, ar;d that in the case of 
species growing "naturally together Ihe potential environment of the individual 
species overlaps the actual environment. In no case is there evidence of 
succession or progression to a climatic climax. 

In the system of classification of communities proposed, the unit (the 
association) is based upon constant association oT dominant species in which 
the term dominant, refers not only to tree species, but io characteristic species 
of other layers. The concept of edaphie comple.w firsf: proposed as a unit of 
convenience, is shown by practical experience to be especially valuable in view 
of the complex soil mosaic consequent on the previous history of the area 


1. Au analysis is made of some of the factors affecting the development 
of the Australian flora in the lighl of an improved knowledge of the geological, 
pcdologicat and ccnlogicnl pattern of Australia, and in relation to the more recent 
paleohotanical literature. 

2. Evidence for a .mid-Recent period of aridity in Australia is brought 
together and summnrised, The significance of this desiccation on the HishihutJon 
pattern of species and communities in South Australia is discussed in some detail. 
The presence of a large pre-arid relic, clement in the flora k postulated, and reasons 
given to explain the presence of a large number of ''species complexes/' 

3. A number of Australian plant geographical prohlcrns arc considered, Tbesc 
include tbe origin of the Australian element, and the high degree of endemistu in 
the south -west of Western Australia. 

4. ideological concepts are discussed in the light ot ihe time factor involved. 
The bases underlying systems of classification are discussed and ecological units 
are defined. 

<*> C, G. Stephen*. Bull. 206, O'.J.R. (Aukm. in pre?*. 



The authors wish to acknowledge information on the distribution of certain 
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VOL. 71 PART 2 DECEMBER 22 1947 





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By Nancy T. Burbidge, M.Sc. 

The publication, in 1934, of Blakely's "Key to the Eucalypts" marked a definite advance in our 
knowledge of the genus, since the book made a very useful companion to Maiden's rather bulky 
"Critical Revision of the Genus Eucalyptus." It was a pity that Blakely's work, which was the result 
of years of patient and intensive labour, appeared under such misnomer, since identification of 
specimens with the aid of his "key" is difficult even when one has acquired a certain familiarity 
with the genus. This is particularly so when dealing with some of the members of such a complex as 
that around E. odorata or E. oleosa. Owing to the number of new species described by Blakely and 
the modification of some of the older ones, the key in Black's Flora of South Australia is no longer 
adequate and a new one is needed. 



By Nascy T. BuumooK, M.Sc. 
Waile Agricultural Research Institute, Adelaide 

fRcad 10 April 10471 

The publication, in 1934, of Blakely's u Key to the Eucalypts" marked a definite 
advance in our knowledge of the gpilWa, since the book made a very useiul com-, 
panion to Maiden's rather bulky v 'Criiic;il Revision of the. Genus Eucalyptus-" 
It was a pity that Blakely's work, which was the result of years of palieni and 
intensive labour, appeared under such misnomer, since identification of specimens 
with the aid of his "key" is difficult even when one has acquired a certaia 
familiarity with the genus. This is particularly ,so when dealing with some of the 
members of such a complex as that around /\. odomta or E. oleosa. Uwing to 
the number of new species described by Blakely and the modification of some of 
the older ones, the key in Black's Flora of South Australia is no longer adequate 
and a new one is needed. 

Since. Blakely's death no botanist has taken over his cucalyptological work, 
which is lu he regretted when the economic and botanical importance of the penus 
is considered. The whole genus is badly in need of integrated research by 
taxonoftiisi*, eeologists and geneticists, and hence this paper merely tries to 
clarify the present confusion so that a key to the South Australian species can 
be constructed. 

The genus Eucalyptus includes, an unusually high percentage of variable and 
unstable species which, for want of a more definite, term, can be called poly- 
morphic, Hybridisation is known to occur in the field, and evidence concerning 
its extern in Tasmania is being accumulated by Brett (5). Under natural condi- 
tions, aud with such slow growing aud long-lived species, il mnst be a long period 
before the results of any cross can be stabilised. It is possible thai complete 
stability is never attained, arid the plants representing various stages rimy be found 
clo*«* together in one locality or scattered throughout a region according lo their 
adaptability lo different habitat conditions- Possibly this explains the existence, of 
some of the species complexes. But, whatever the cause of polymorph}' may be, 
it is the n suiting variants which must be deal with in a key. 

When there is a scries of related forms, the status given to any particular 
variant depends, partly upon the amount of material available for cotupin.son. 
If specimens A., C and E. are found they may all apparently deserve specific 
rank. Add D. and they may be reduced to varieties. Add B. and F. and it 
becomes very difficult to separate them ar all under any reliable cbnuicter. <)n 
the other hand, the larger and more polymorphic a species becomes, the harder 
it is to delineate il sharply bom other related specks. Hence a balance must be 
struck, and since the meaning of the word "species" is, under these circumstances, 
more than usually uncertain, one is forced to take a -somewhat nlibfarian view 
and, seek the most workable arrangement for practical purposes. 

This work is not intended as a complete revision of the local species which 
would require some years of sUidy both in the herbarium and in the field, as vvclt 
as a fuller knowledge of Eucalyprs occurring in other parts of Australia. The 
writer is in full agreement with Ferdinand von Mueller when he states, "to assign 
to each species its proper place involves the study of ail allied congeners, and 
these are often .... dispersed at wide distances in Australia'* (9). At present 
there are large gaps in localities whence specimens have been obtained. Until 
the areas around the Great Australian Right and to the north of the NuHarhnr 
Plain have been studied, relationships with Western Australian species caono! 

Triins. Ror. Soc. JS- Ausi., 71, (2), I Oucrmbcr 1047 


be solved. Central Australia is still, bolamcally speaking, terra iucoguita and 
much remains to be karat concerning ihe southern species which extend north and 
the tropical forms which occur over the hmver o[ Sooth Australia. In the eas:, 
affinities with species in New South Wales and Victoria are better understood, 
bul knowledge is far from complete 


in his preface Blakely states thaL the original intention of Maiden was to 
design a key based on the nature of the cotyledonary and juvenile foliage- Owing 
to incomplete data this proved impossible and Blakely used ihe anther type in his 
primary division of the genus, The shape and number of juvenile leaves is likely 
to be of real importance when relationships are being traced, but in a key such a 
character would be difficult to employ since the necessary material is not always 
available to the collector. In basing his key on the anthers Blakely followed 
Ben than i (li, but. unfortunately, great familiarity with his herbarium material 
and a certain lack of held experience led to subdivision into a large number of 
groups based on slight differences in shape. The result is confusing both to the 
novice and to the experienced botanist. The original plan as designed by Bcnihuru 
was reasonably sound, though until adequate anther material of all species is avail- 
able the value of this character cannot be fully estimated. Blakely '$ subsections 
are so difficult to recognise that anthers have been omitted from this key, except 
where several features of the organ can be contracted with corresponding 
characters in other specie?. 

Bentham's secondary divisions were largely based on the character of the 
fruit, and in the preparation of this paper more -and more reliance came to be 
placed on this feature, particularly on the nature of the disc and the type of the 
valves. Correlation between anther type and fruit character was imperfect, though 
it exists to some extent-. This issue is probably confused because oi the number 
of line& of development which are represented among the specie*. 

The top of the fruit is formed of four zones. The outer rim represents the 
abscission line left by the fallen operculum and is known as the calycinv viag f the 
next is the staminal rhuj, then the so called disc whose ontogeny has not yet been 
described. It is not certain that this term has been consistently employed by 
butanists. Inside the disc is the top of the ovary which ruptures into the .r'tf/rc*. 
The variations produced in these zones in different species are considerable, The 
calycine ring is rarely important, though it is wide and conspicuous in the flowers 
and fruits of E. pyrifonuis and its affinities. This tact has been employed in the 
key In some cases the staminal ring is a narrow elevated ring in the rlovver, but 
it usually becomes inconspicuous in the mi:i, In others, e.g., E. icuctKvykm, tltp 
ring is comparatively wide, but thin and projecting over the* disc. In this species 
the ring either falls away during maturation of the fruit or it remain* partial V 
attached on one side of the frnir In E. httcrtexta, and also in E. calcicullrix, 
which lies between JjL teitcoxylon and E. odorata there may also be a distinct 
staminal ring in. the maturing fruit, but usually it is not deciduous und it may 
break up into a number oF small pieces attached around the rim of the fruit. 

In fresh flowers and young fruits the gfeci is more or less continuous with the 
lop of Ihe ovurw As the fruit matures or, in collected specimens, dries the dis- 
tinction between the two pails becomes increasingly marked until, with the open- 
ing of the valves, they appear quite unrelated. The disc itself may be broad as 
in £. Baxteri, a narrow ridge as in E_ tncntssata, a ring as in jfiL oleosa or E t Icpto- 
phylla. or it may be little more, than a lining to the" lop of the tube when the 
capsule is sunken as in E. fasckulosa rtr E. c'orota. When broad the disc mav 
be domed, i.e., convex as in G, camahtnfeusis and E. viviuwtis; rial as in K Htrr'u 
ot enncave, .'.c v oblique a* in E, obliqua and E. leuvoxylon. 


The valves may be short and triangular as In H. canidhiulensis in which case 
they represeut Lhe top of the ovary, or They may have sharp points formed by the 
rupturing of the persistent base of the style as in /?. incrussata and E. tlumosa, or 
the sharp points may he long and coherent at the apex clue to the persistence oi the 
entire style as in E. oleosa, in the larter case the valves are brittle and may break 
off short, confusing the inexperienced student. 

Inst which type can be regarded as the most primitive is a question demand- 
ing a wider and more extensive study and eorrelalion with the affinities demon- 
strated by such characters as juvenile' foliage and anther shape. Owing to South 
Australians central position, geographically, the local Eucalypts include species 
representing a number of different lines of development. Consequently much 
speculation here concerning phylogcnv within the genus would be out of place. 

In specimens hearing" immature ft nit the valves open during drying and 
release the seed Such capsuk-s may appear fully developed and will give a false 
impression of the size normal for the species. In plants growing in arid habitats 
a lack of soil moisture or a sudden hot spell at the crucial time may induce a 
similar dehiscence of immature fruit. Another probable cause of subnormal size 
and shape in fruit, apart from iusect or mechanical damage, is when no seeds 
mature. This is suspected when /:". Morrisii^ which is supposed to have a high 
convex disc, hears fruits with flat ones (pi. u hfif. 4b-c). 

The word ''striate" has caused some confusion in the literature. Clakely 
used it rather freely whenever the bud or fruit was slightly marked by longitudinal 
lines. Others have allowed the term to cover fruits which Blakely would have 
called ribbed. In this paper the word is u&ed when longitudinal ridges or lines 
are apparent owing to the shrinkage of non-sclempd tissues, e.g., E. incrassuta 
var. costata, while "ribbed" has been used when the ridges arc deep enough to have 
been evident when the specimen was fresh, e.g.. E. fiyriformis and E. in eras so i 'a 
war. angidasxi. In a few cases it is difficult to draw a line between the two condi- 
tions, but in general the above use of the terms can be quite consistent. 

With reference to distribution, it may be stated here that Blakely > lists oT 
localities were compiled from the material in the Sydney Herbarium. They are, 
therefore, incomplete in some cases,, E, dacopkora, and incorrect in others. 
e.g., E. goniocalyx. The latter reference was based on a specimen which is, in tbc 
writer's opinion, E. elaenphora. 

Short descriptions of the species have been included to make the use of the 
kev easier. Field details such as bark, wood colouration and juvenile foliage have 
been omitted. The writer cannot claim a wide knowledge of the first two. and 
ail have been covered in Bhkely's descriptions. 

Key to the Species 

(N.B. — This key is based on appearance of ^nmrnens whtn dry. Ribbing 
01 wrinkling of buds and fruits is always less evident in fresh material.) 

A. Staminal ri#g lobtd, -Buds telragonrnjs. Leave* oyposiic -- 1. trwtcsmiouUs 

AA. Stamina! ring net lotted 

B. Operculum thick, leaving abroad rim -outside the stamina! ring. 
Disc with an erect rJtfee around the partially sunken valves. 
C. Bud* ricclicHLiu ■, calyx in)>c rilibed, 

D. Fruit more Oian 4 cm. in diameter ... 2 pynformis 

DD. Fruit less than 4 cm. fa diaiiK-ter . .,, 3. pttrhypkylh 

CC. Kuds sessile, catyjc smooth. 

D. Kudo giancoub .... ... ... 4. o.rvtui-lra 

DD. Buds not ^,-lsui'mu ... .... ., ... 5, Oldfnidii 

BD. Operculum not thick. 

C. Disc broad and convex or flat arounrf tho shod broad -vaUe;- 
T).'vq either markedly \ xsrrt .inn contributing to the ItupjUJ 
01* thf- tfutt or in the orifi-.L-. 



Peduncles and pedicels slender, 

Operculum rostrate or 

EE. Peduncles and also pedicels when present not slender, 
usually short and stout. 
F, Malices or small stunted trees. 
G. Species occurring in arid interior. 

H, Operculum conical to rostrate. Fruit almost or 
quite sessile., 

I. Operculum acuminate to beaked. Bud glaucous .. 
IT. Operculum Bnris not glaucous .... 

HH. Operculum hemispherical. Fruit pedicellate 
GG. Coastal or subeoastal species. 
IT. Operculum hemispherical or hemispherical-conical. 

Buds and fruits pedicellate 

HH. Operculum conical to rostrate. Buds and fruits 

very shortly pedicellate or sessile 

FF. Trees occurring in higher rainfall areas. 

G. Fruit about 10 x 10 mm., markedly sessile (except 
var.). usually in globose clusters. Operculum hemi- 
spherical or short and very obtuse , 

GG. Fruit smalVr than above. Operculum conical. 
TT. Juvenile leaves ovate-orbicular, glaucous. Buds and 

fruits more or less glaucous 

HH. Juvenile foliage broad or narrow lanceolate, pale 
green. Buds and fruits not glaucous. 
T. Umbels with three flowers 

II. Umbels with three or more flowers occurring" on 
same branch. 

J. Umbels on leafless branches or racemes 

JJ. Umbels all axillary 

DD. Valves enclosed and more or less concealed by the disc. 
E. Operculum conical, about as long as the tube. Venation 

semi-longitudinal, Bartr rough 

EE. Operculum shorter than tube, hemispherical. Venation 

longitudinal. Bark smooth 

CC. Disc narrow and forming a ring or rim on the top of the fruit 
or, if broad, concave and extending obliquely into the 
capsule or forming a lining inside the top of the fruit. 
D. Disc forming a ridofc or concave and oblique. 

E. Operculum hemispherical - apicu!ate«. Disc strongly 
oblique t '__ 

EK. _ Operculum conical or rostrate (except H. ccncintia). 

G. Slaininat ring 1 obscure or if evident on fruit then 
not broad and extending over the disc. 
IT. Flowers in axillary umbels, 
I Stamens all fertile Outer ones not markedly- 
longer than inner. Buds without conspicuous 
oil glands. 
J. Ambers versatile, opening widely to ba&e with 
cells parallel and side bv side, with a con- 
spicuous gland on back. 
K. Trees in high rainfall country. 

L. Buds not angular at base. Fruit turbinate 

6. camaldulensis 
Sc var. obkisa 


Buds angular 
Ala 1 lees 

it base. Fruit cylindrical 

Buds and fruits sessile on very 

id peduncles, bicosttUp - 
ribbed. Top of ovary not ridged between 

, short but 
bicostate or un- 

not flattened peduncle 

ribbed. Top of ovary 

valves when immature. Valves" in orifice . 

4. pxytnilra 

7. Morrisii 

8. Ewartiana 

9. re mo fa 
10. tlk'&rsi 'folia 

lir Ba-vtcri &var. 

12. rubida 

13. z'imiiialis 

13a. t>imiinztis var. 

13b. vimbiaHx var. 


14. vitrea 

15. pauci flora 

16. obliqua & var. 


17. ovafa & var. 
ffrandi flora 

18. chwophora 

19. cosmophylla 



LL. Buds and fruits pedicellate, or if sessile 
and bicostate then with flattened peduncles. 
Top of ovary usually ridged between valves 
in immature fruits. 
M. Valves deeply enclosed. Umbels defle.xed 
MM. Valves in or near orifice. Umbels 
N. Valves broad and triangular with short 
points. Fruit 10 x 10 mm. or larger. 
O. Fruit smooth. Peduncle terete .... 
OO. Fruit striate. Peduncle terete or 
narrow but flattened .. 

OOO. Fruit coarsely ribbed. Peduncle 
broad, flattened .... 

NN. Valves narrow and more or less sub- 
ulate. Fruit smaller than above. 
O. Buds and fruits short and fat, the 
latter broader than long and so sessile 
as to be almost imbedded in the end 
of the flattened peduncle 
OO. Buds and fruits not as above. 

P, Fruits sessile, striate, on thick 

PP. Fruits pedicellate, sometimes only 
shortly so. 
Q. Calyx tube 2-4-ribbed and pass- 
ing into a flat pedicel. Peduncle 
flattened. Operculum hemi- 
spherical-conical, obtuse 
QQ. Calyx tube not 2-4-ribbed. 
R. Peduncles and pedicels not 
slender. Operculum shorter 
than tube, 
S. Buds smooth or very 
minutely striate. Peduncle 

terete .... 

SS. Buds with radiately ribbed 
or striate opercula. Peduncle 
angular or flattened 
RR. Peduncles and pedicels slen- 
S. Operculum as long as the 
cupular calyx tube, obtuse ... 

SS. Operculum shorter than 
tube, hemispherical and flat- 
. Anthers more or less adnatc to the top of the 
filaments; dehiscing by pores, slits or by broad 
cells opening back to back. Valves narrow and 
K. Filaments sharply re-flexed in bud and with 
a bend when flowering-. 

L. Buds and flowers pedicellate 

LL. Buds and flowers sessile 

KK. Filaments crumpled in bud, with or without 
some reflexed, not bent when flowering. 
L. Buds and fruits sessile. Leaves erect 
LL. Buds and fruits pedicellate. Leaves 
M. Fruit globose to sub-urceolate. Opercu- 
lum not w T ider than tube at junction. 

20. pimpiniana 

21. incrassata 

21a incrassata var. 

21b. incrassata 
var. anaulosa 

22. conglobaia 
2 j. anceps 

24, rugosa 

25. dumosa 

26. pileata 

27. brachycalyx & 
var. c hind oo 

28. concinna 

29, h'plophylla 

30. uncinata 

31. cncorifolia 


N. Leaves opposite, sessile or almost so. 
Plant usually very glaucous. Operculum 
rostrate , r 

KN. Leaves alternate, petio!ate. 

O. Operculum with the same surface 
texture as the tube. 
P. Operculum conical. 
Q. Leaves shining and glossy 
QQ. Leaves, buds and fruits glau- 
cous .... 

PP. Operculum rostrate. Leaves dull 
and more or less glaucous .... 

OO. Operculum very obtuse or flat- 
topped, the surface wrinkled and 
usually a different colour from that 
of the smouth tube 

MM. Fruit urceolate. Operculum smooth, 

wider than tube at junction 

IL Outer stamens sterile and longer than inner. 
Buds with translucent oil glands. 
J. Buds and fruits quadrangular 

JJ. Buds and fruits not quadrangular 

HH. Umbels paniculate, sometimes simple axillary 
umbels also present. 

I. Buds 7x5 mm., valves of fruit narrow, enclosed 
in orifice - 

II. Buds 4x3 mm., valves broad and exsert 

GG. Stamina! ring broad (up to 2 mm.) and conspicuous- 
after flowering, more or less deciduous. 
H, Three flowers per umbel. 

I. Fruit smooth. 

J. Fruit 7-15 mm. long 

JJ. Fruit more than 15 mm. long , 

II. Fruit subangular, 4-6-ribbcd 

HH. Umbels with three or more flowers. Fruit 
6-7 mm. long .... .... . ., 

DD. Disc very inconspicuous and lining the rim of the calyx 
E. Flowers in axillary umbels which may become clustered 
or terminal due to lack or loss of leaves. 
F. Leaves more than 0-8 cm. wide. 

G. Buds crumpled or wrinkled like a withered apple. 
Leaves lanceolate to broad lanceolate, tbe marginal 
nerve distant and the leaves often tri-ncrved at the 
base . .. ,_ 

GG. Buds more or les^ angular with angular pedicels, 
not wrinkled like a withered apple. 
H. Fruits more than 4-5 x 4-5 mm. 

I. Buds sessile or on very short thick pedicels. 
Leaver lanceolate or broad-lanceolate, rather 
thick. Mallee .... 

II. Buds pedicellate, often markedly so. Leaves 
narrow lanceolate or, in tree forms, broad 

HFf . Fruits 4-5 x 4-5 ram. Buds very shortly pedicel- 
late, slightly glaucuos. Leaves narrow lanceolate .. 

32. Gillii & var. 

33. oleosa 

33a. oleosa van 

33b. oleosa var. 

33c. oleosa var. 

34. Flocktoniae 

35. calycogona & 

var. StaSJordil 
36. gracilis Si 
var. crecta 

37, intertexta 8c 
var. fruticosa 

38. coofabah 

39. leucoxylon 
39a. leucoxylon 

var. mcacracctrpa 
39b- leucoxylon 
var. angtttala 

39c. leucoxylon 
var. paupe-t ila 

40. calcicullrix 

41. iMnsdoivneana 

42. odorcUa 

42a. odorata var. 


FT Leaves narrow linear. leg* th*n 0*8 cm. wide 

FF. Flowers in paniculate umbels. 
F. Leaves alternate. 

G. Fruit smooth or with one or two striatums, not 
H. Operculum as wide as the top of tube. 
I. TCuds and fruits pedicellate. Trees. 
J. Operculum conical, 

K. Fruit larger than 5-7 x 4-5 mm. 
L. Leaves, buds and fruits glaucous .... 

LL, Leaves, huds and fruits not glaucous .. 
KK. Fruit 5-7 x 4-5 mm. (sometimes almost 


JJj Operculum hemispherical 

IL* Buds and fruit? sessile or very shortly pedicellate 
(see also hemiphhio var, mhroairpa above,). 
J. Operculum conical. Fruit subanKular, !1 P ^ 

10 x 9 mm. Leaves lanceolate .... 
J J. Operculum hemispherical. Fruit 3-S x 3~$ mm. 
Leaves broadly lanceolate 
FIH. Operculum conical, narrower than top of tuhe .... 
GG. Fruit tirceolate, costate. Operculum hcmispherical- 

apiculate , ■ 

FF. Leaves opposite - — * ;■ 


43. I'it'idis 

44a. hcv'jphk'ia 
var. o&bcns 

44. hcmil'hfokt 

44h. hemiphkm 
var. vihrocarpa 

45. larytjlorens 

A<t. Bghriana 

47. fasciriiiosa 

48. clailocalyx 

49. yumuphyUa 

1. Eucalyptus euuesmtoiues F. Mueli. in Fragm., ii, 35 (1860). 

Dwarf mallee or small tree. Mature leaves opposite, shortly petiolate, 
lanceolate, stibglaucous, 4-6 x 1-2*5 cm. Umbels axillary, three-flowered, 
peduncles terete. Buds clavate, teiragonous, obtuse, shortly pedicellate, 
6x4 mm. Operculum hemispherical shorter than tube. Stamens in four 
clusters alternating with small calyx teeth. Fruit pedicellate, campanulatc to 
ureeolaie, smooth or slightly angled, 14 x 10 mm. (afler Blakely)- 

This species, recorded for the Birksgate and Blyth Ranges in the extreme 
north-west of the State, has no close affinity with any other South Australian 

2. Eucalyptus ryRu-omns Turc?. in Bull. Soc. Nat Mose., 22, pt: ii, 22 


Mallee with straggling habit. Mature leaves alternate, pelio?atc, broadly 
lanceolate, 6-15 x 2-4 cm., surface dull \hitbels axillary, 3-flowered, peduncles 
defiexed, thick and short. Duds ovoid or pyriform, shortly pedicellate,, 5-6 x 
3-4 em, Operculum conical with a rostrate apex, very ihirk but narrowing to 
about 4 mm. thickness at the point of attachment to the calyx tube, striate, twice 
as long as tube. Tube wide and shallow with 7-9 coarse ribs and a few smaller 
ones. $!amcns red or yellowish. Fruit broadly turbinate, more than half rhe 
height being due to the high disc. Staminal ring present as a line separating 
the calycine ring and disc, the latter erect and with a distinct ridge around the 
broad, thick, deltoid, partially sunken valves. Fruit 4x7 cm. (pi. i, fig. 2). 

This species, with E, pachyphylla and E. Oldfieldn, is distinguished by the 
thickness of the operculum and by the ridded disc which forms a small crater- 
like valley around the valves. These characters are less distinct in E. Oldfieldii, 
which forms a link with E. oxymitra, E. Ewirliana and F., Mnrrisu. 

It is known to occur along the transcontinental railway from Tareoola 


3. EucALVrrus F. Muell. in Jour. Linn. Soc, 3, 98 (3859). 

Malice. Mature leaves alternate, petiolate. ovate or broad to narrow 
lanceolate, 6-10 x 2-5 cm. Umbels 2-3-flowered, axillary, peduncles terete, 
spreading not deflexed, 6-10 mm. long. Buds shortly pedicellate, ovoid-rostrate, 
25 x 20 nun. Operculum conical-rostrate, striate to ribbed, about 2-3 mm. thick 
at point of attachment, 1-1 J times as long as broad. Tube wide and shallow with 
3-5 sharp hut narrow ribs and with smaller ones hetween. Frtitfs a& in E. pyri- 
formis but smaller, 15-20 x 20-30 mm., ribs more rounded in older fruits (pi. i, 
% 3). 

Description oF the bud was taken from Maiden (Ctit Rev., pi. 75. 
fig. 5-6). This species was collected by Professor J. B. Cleland at the junction 
Oi Boundary and Fraser Creeks on the border of South Australia and Northern 

4 HccAr.YPTUs oxymjtka Blakelv in Traus, Rov. Soc S, Aust., 60, 155 

Glaucous mallee. Mature leaves alternate, petiolate,, broadly lanceolate, 
thick, 5-7 x 2-4 cm, Umbels axillary, 3-7-rlovvered, peduncles short. Buds 
shortly pedicellate, ovoid or globular with a hooked-rostrate apex 10-12x7-8 mm., 
glaucous Operculum hemispherical-rostrate Or conical-rostrate, 2-5 tunes as 
long as wide shallow tube. pedicellate, glaucous, subglobosc, 11 x 11 mm. 
disc broad, semiconieal, smaller than calycine portion (after Blakely) (pi. h 
fig. 1) 

Fruiting specimens used by Blakely have not been seen by the writer. The 
affinities of this species and also the identity of S. A. White's Everard Ran^t 
specimens (see below) depends on whether the disc in this species is ridged 
around the valves or not. Its occurrence in this State is still in doubt. 

5. Eucalyptus OLwi&wm R Muell. in Fragm., ii, 37 (I860), 

Mallee. Mature leaves alternate. pejtiolate, narrow to broad lanceolate, 
7-13 x i*5-3 cm. Umbels axillary, .3-flowered, peduncles thick, sometimes very 
shorr. Buds globular to ovoid-rostrate, almost or quite sessile, 20 x 10-15 mm. 
Operculum conical or rostrate, thick, longer than tube. Tube smooth, wide and 
shallow. Fruit hemispherical to turbinate, 11 x 15 mm.; disc portion nearly as 
large as base, ridge and hollow around valves less evident than in E, pyriformis 
and K parftyphyfla (mostly after Blakely). 

There is some doubt as to whether this species actually occurs in South 
Australia. ftJick (Fl. S. Aust.. 418) mentions a specimen from Missionary 
Plains, MaeDonnell Range under E. Oldfu-ldii which has since been describe/.! 
by Ulakely as hi, oxymitrz (see above), The specimen consists of buds and 
leaves without fruits. Of S. A. White's material mentioned by Black (/.<".)» 
both have leaves and fruits but no buds. Tn his MaeDonnell Range specimen 
the disc is ridged though rather smaller than is illustrated tor E. Drumnwndii by 
Maiden (Crit. Rev., pi. 74, rig. 4 b). while in the Everard Range material 
the fruits have a convex disc, reminiscent of that in E. Ewartmna. Pending 
collection south ot the border ihe recording of E. Oldfietdli for this State must 
remain douhtiuh 

6. Eucalyptus cam ALuuuiNsis Dchn in Cat. PI. Hort. Camald. ed. 2. 20 

(1832). E. rostrafa Schlecht in Linnaea, 20, 655 (1847), 

Medium to large-sized tree. Mature leaves alternate, petiolaie, lanceolate, 

5-11 x 2-4 cm. Umbels axillary, 5-10-flowered, peduncles and pedicels slender 

and Lerele. Bads ovoid or diamond-shaped, acute, pedicellate 6-10 x 3-5 nun. 

Operculum conical and acute or rostrate, longer than the wide shallow tube. 


Print hemispherical or broadly cupular with tltc erect valves at least as long as 
the wide base, 6-8 x 5-6 mm,, the valves broadly triangular and more or less 

This widespread species occurs throughout the State. It is associated with 
watercourses and creeks. 

7. Eucalyptus Mouristi R. T, Baker in Proc. Linn, Soc, N.S.W., 25, 312 

Malice or small tree. Mature leaves alternate, petiolate, linear-lanceolate to 
lanceolate, 6-12 x 12 cm. Umbels axillary 3-6-flowered, peduncles semircrctc or 
Terete, 5-10 mm. Joilg, Buds ovoid, twiiscssile on short thick pedicel? which 
may he slightly angular ad they pass into the calyx tube, 7-10 x 4-6 nnn„ with. 
copious dark oil glands. Operculum conical, very obtuse, l£-2 times &a long as 
tube. Tube obeonical, about as long as broad, smooth or with one or two small 
ridges: Fruit truncatc-turbinatc or ovoid according to the convexity of the wide 
disc, valves short and broadly triangular, exsert at the top of the disc (pi. i> 

fig' 4 )- .... 

With regard to the variation in the development of the disc. Maiden (Cnt. 

Key., pi. 135, fig. 6) figures the disc as continuous with the valves. This is, 
always more marked in fresh material than in dried. Maiden's fig. 6 was 
apparently taken from R. T\ taker's original plate, but neither author states 
whether the specimen used was fresh or dry. Tn material collected by the writer 
an excellent match with Maiden's fig. 6a and 6b has been obtained. In most of 
the South Australian specimens the buds and fruits are smaller than the dimen- 
sions given for New South Wales material. 

In this State the species is found in the Flinders Ranges, from Quoru 

8. Eucalyptus EwartiAna Maiden in Jour. Roy. Soc. N.S.W.. 53, 111 


Malice. Mature leaves alternate, petiolate, lanceolate, 5-9 x 1*5-2*5 cm., 
surface dull, Umbels axillary, 3-7-fl owc-rcd, peduncles terete, up to 2 cm. long. 
Buds pedicellate, clavate or globular, 8-10 x 6-8 mm. Operculum very obtuse 1 , 
being hemispherical or basally truncated-spherical, thick, 1-H times as long as 
lube, smooth. Tube wide and shallow, half as long* ■as wide. Fruit pedicellate, 
plobose or clavate due to the very convex disc which provides neatly half the 
length, valves deltoid and very exsert, 8-10 x 8-12 mm. (pi. i, tig, &J, 

Despite the fact that Maiden (Crit. Rev., 44, 120) stated that the anthers 
placed this species among the Macrantherae, Blakely included it with the 
Platyantherae, Blakely records this species for South Australia, but gives- a 
Central Australian locality — Missionary Plains. MacDonnell Range. Klack (3) 
states that the valves have long points (tip to 4 mm. long), which break off early. 
These have not been seen by the. writer, 

9. Eucalyptus rkmota Blakely in Key to Euc, 197 (1934). 

Malice. Mature leaves alternate, petiolate, lanceolate, to broadly lanceolate. 
slightly falcate resulting in better development of the base on one side than on the 
other, coriaceous, 6-12 x l'5-3 cm. Umbels axillary. 3-8-rlowered, peduncles semi- 
terete, 5-11 mm. long. Buds pedicellate, clavate. 4-6 x 4-5 mm. excluding the sub- 
angular pedicels. Operculum hemispherical or henuspherical-apiculate, about as 
long as the tube. Tube obconical. passing into the pedicel to form a long narrow 
funnel, surface wrinkled in dried material. Fruit pedicellate, pyriform. slightly 
wrinkled when dried, flat-topped, 6-9 x 5-8 mm., disc slightly convex and rounded 
lo very slightly concave but usually flat, about 1 mm. bruad; valves xhorr, in 
orifice or almost covered by disc (pi. ii, fig. 1). 


Blakely placed this species near E. Cansideniana and E. Sicberianct, which 
both belong to eastern New South Wales, Victoria and Tasmania, However, to 
the writer the relationship with E. dtversi folia, which occurs in the same localities 
as E. reivota, is far mure marked, especially in the fruits. The rmds and siiglitly 
asymmetrical leaves suggest a link with E. obUqna. E. remola is known as a 
useful indicator plant for a soil type on Kangaroo Island, which is, so tar, the 
-only area in which the species has heen found. 

10. Eucalyptus diykksjkoua Bonplaud. in PI. JnrrJ. Malm., 35, t. 13 (1813). 

E.mntaUfolia F. Muell. in Trnns. Vict. Jnst., 1, 35 (1855). 

Large Mallee. Mature leaves alternate, petiolate, linear-lauceolate to broudlv 
lanceolate. Umbels axillary, 3-6-fIovvered, peduncles scmiterofe 4-7 nun. long. 
Buds shortly pedicellate or subsessile, shape of two cotics attached hasc to base, 
smooth or very sh'ghtly wrinkled when dry, 7-10 x 4-6 mm. Operculum conical 
or conical-rostrate, about the same size and shape as the combined tube and 
pedicai, 4-6 mm. Joiir^. Tube obconieal, 3-4 mm. long. Emit sessile or very 
shortly pedicellate, hemispherical to campamdate or obconieal, 8-12 x 10-15 mm., 
disc 1*5-2 mm. broad, slightly convex and rounded or Sat; valves broad and 
triangular in orifice and slightly more exsert than in E. r?mata_, so that the short 
points protrude above the disc (pi. ii, §& 3). 

When buds are lacking it is not easy to distinguish between IL dlversifolin 
arid Ii. renioia, but the fruits of the former are larger, have a wider disc and are 
always more shortly pedicellate than those of the latter. 

This species occurs as a coastal or subcoastal malice from the west coast ot 
Eyre Peninsula to the Victorian border, including Kangaroo Island. 

11. Eucalyptus Baxteri (Lenth.) Maiden and T.Iakely In Crit. Rev., 70, 451 

(1928) E. satttaiifalht var. ( ?) Baxteri Benth. in EL Austral.. 3, 
207 (1866), 
Small to medium-sized tree, Mature Icai^es alternate, petiolate, ovate to 
bmadly lanceolate, asymmetrical at base. Umbels axillary, 5-9-flovvered, peduncles 
stout 4-6 niru. long. Buds obovoid on short thick pedicels. 4-5 x 3-4 mm. 
Operculum hemispherical, slightly shorter than the obconieal or hemispherical 
tube. Fruit sessile, subglobular due to the convex disc which is 2-3 mm. wide, 
9-12 x 10-13 mm., arranged in tightly packed globose clusters on the short stout 
peduncles, valves incurved, in orifice -or slightly exsert (pi. iii, fig. 1). 

11a. E. Baxtkri var. pediceu.ata Maiden and Klakcly in Crit. Rev., 70, 457 

Differs from the above in that the fruiis are shortly pcdicellaie. The opercula 
are vcrtucose. 

The species occurs in the Mount Lofty Kange and southwards to the South- 
East and the Victorian border, also Kangaroo Island. The variety as kuown to 
occur in the South-East and may be found in other parts of the above districts. 

12. Eucalyptus rtjbiua Deane and Maiden in Eroc. Linn. 5oe. N.S.W., 21, 150 


Medium to large-sized tree. Mature leaves alternate, petiolate, lanceolate. 
U}(ihcfs axillary or lateral on leafless portions of the branchlels. 3-ilowered, 
peduncles semitcrete or slightly flattened. Buds shortly pedicellate or almost 
sessile, ovoid or cylindrical, 5-7 x 3-5 nim. Operculum conical, obtuse. Tube 
cylindrical, slightly longer than or equal to the operculum. Emit sessile or shortly 
pedicellate, obovoid or turbinate, 5-6 x 6-7 nun, ; disc concave, calves deltoid and 


The South Australian material seen was less glaucous than a* described by 
Blakcly. It appears to lie baween IL eloeophora and E. viminalis among our 
species. It occurs in the Mount Lofty Range. 

1.1 Eucalyptus viminalis Lab ill in Nov. Ho1L, h\ 12, t. 15 L (1SQ6). 

Targe tree. Mai are /eaves alternate, petiolate, linear-lanceolate to lanceolate, 
10-20 x 1-2-3 cm. Umbels axillary, normally 3-flowercd, peduncles suban^ular, 
3-6 mm, long. Bttds obovoid wilh pointed apex and short usually thick pedicels. 
6-10 x 3-5 mm. Operculum conical to conical-rostrate, equal to or slightly longer 
than the obconical tube, at its broadest somewhat wider than tube, fruit sessile 
or very shortly pedicellate, turbinate to subgVihnlar or pyriform, 6-8 x 6-9 mm. 
(including valves); disc convex, contributing to the length of the Imit, about 
1 mm. wide, valves exsert and erect, short and broad r 

Occurs hi the Mount Lofty Range and southwards lo Mounr tJambiYr atid 
the Victorian border. 

13a- Eucalyptus vimikaus var. kAchmosa (F. MuellT Biakely in Key tn Fnc., 
162 (1934). 

In this variety the umbels arc arranged on short leafless branches. Elakely 
states that the umbels are 3-flowercd, hut South Australian specimens agreeing 
excellently in other respects with the figure (Crit. Rev., pi. 118, fig. 9) have multi- 
flowered umbels. There is no reference in the text of the Critical Revision to this 
variety, and Blakely's note appears to be the only description. The variety occurs 
in the South-East of the Stale, 

13b, Eucm.yftus var. HurtERTANA (Naudin) N, T, Rutbidge stal, 
■nov.; E. HunKuiArtA Naudin in Second Mem.,, 42 (189f). 

iB. liubcrimm was separated from E. z'lminalis by Rlakely. though Maiden 
(Crit. Rev., 28, 173) considered it merely a synonym and remarked that it was 
always possible to find multiflowered umbels on trees with mostly 3-flowercd ones. 
£. Huheriana differs in timber value, rougher bark, form of growth and in the 
multifkrwered umbels. Recent specimens, collected near Adelaide, showed some 
branchlets wilh 3-flowered umbels and some with four or more flowers,. This 
supports Maiden's observation and there seems to he no character which can be 
used for differentiation at the specific level and I he form is herein given varietal 
rank (pi »ii, fig. 2). Naudin';-, species was based on a -single tree growing under 
cultivation at Nice. This is a praclice which is open to criticism. Many Jvuca- 
lypts produce an atypical growth under horticultural conditions, and also there is 
no evidence concerning the origin or purity of the seed used. 

Distribution is the same as in R. vivu'ualis, also on Kangaroo Island 

14. Eucalyptus vitkea R. JL Faker in Froc. Finn. Soc. N.S.W.. 25, 303 (1900). 

Medium to large-sized tree. Mature leaves alternate, petiolate, narrow to 
broad lanceolate, the venation very oblique and approaching sermlongiiudinal. the 
main nerve some distance from the edge and frequently there arc minor nerves 
outside, 10-15 x 1*5-2 cm. Umbels axillary, singly or in pairs or in short axillary 
panicles. 7-12-flowered, peduncles terctt: or semitcrcte, 7-10 mm, long. Buds on 
relatively stout pedicels which pass into the tube without marked change, 
clavate, 5-6 x 4-5 mm. Operculum hemispherical-apiculate or conical-acute, as 
long as tube. Tube obconical or cylindrical. Fruit pyriform, shortly pedicellate, 
5-6 x 6-7 mm. j disc 1-2 mm. broad, convex or almost flat, covering the short 
enclosed valves. 

In the south of the State. Specimens from Waitpinga, Kalangadoo, Mount 
Oamhier and other South-East localities have been seen, 


15. Eucalyptus pacciflora Sieber in Spreng, Syst. Cur. Post, iv, 195 (1827). 
Small to large tree. Mature leaves alle.rnate, pctiolate, lanceolate, venation 

longitudinal or almost .so, 8-14 x 1*3-3 cm. Umbels- axillary, 5-12-flowercd, 
usually about 9, peduncles thick, terete or compressed 7-15 mm. long. Bud* 
clavate, pedicellate 5-7 x 4-5 mm, Operculum hemispherical, obtuse, shorter than 
tube. Fruit shortly pedicellate, globose to pyriform 7-10 x 5-8 mm.: disc fiat bui 
rather prominent, extending over the enclosed valves (after Rlakely). 

This is a rare specievS in South Australia. No specimens were available 
during the preparation of the above description. It is reported to occur m tilt* 
Mount Gambicr district. 

16. Eucalyptus obliqua L'Herit. in Sert. Aug., 18, 20 (1788). 

Large tree. Mature leaves alternate, pctiolate, lanceolate, asymmetrical at 
base and slightly falcate. Umbels axillary or sometimes arranged in short 
racemes or panicles, peduncles terete or scmitercte, 7-20 mm. long. Buds on long, 
relatively thick pedicels which pass without sharp increase in size into the calyx 
tube, 1CM2 x 4-5 mm. with pedicel. Operculum hemispherical-apiculate, slightly 
wider than tube at junction, shorter than obconical tube. Fruit pyriform or 
ovoid, 7-9 x 7-9 mm.; disc oblique, i.e., concavc\ covering the short enclosed 
valves. In some South Australian specimens the disc is almost flat. 

It occurs from the Mount Lofty Range near Adelaide to the South -East 
and is also found on Kangaroo Island. 
16a. Eucalyptus obliqua var. megacarjm. Rlakely in Key to Euc, 194 (1934). 

This can be distinguished from the above by its coarser buds and larger 
fruits, 10-12 x 10-12 mm., which are more definitely constricted at the orifice 
than in the typical form (pi. ii, fig. .2). 

The type material of the variety, which came from Milhccnt, has been seen 
at Sydney and in the collection of Mr. J. M. Rlack. In other specimens, from 
Eight Mile Creek, in the Waite Institute Tlerhaiium, the buds are often clavate 
and swollen, a malformation apparently due lo some gall-forming inject. 

17. Eucalyptus, ovata Labilf, in Nov., Hull, pi. h\ 153 (1806). 

Large tree. Mature leaves alternate, pctiolate, lanceolate to ovate-lanceolate 
or ovate, 7-14 x 2-4 cm. Umbels axillary or on leafless portion of the branchlets, 
4-8-dowered, peduncles terete or semiteretc, 5-10 mm. long. Buds pedicellate, 
smooth, obconical-rostratc or clavate or like two cones attached base to base. 
7-10 x 4-6 mm. Operculum conical, shortly acute or rostrate, shorter than or 
almost equal to the tube, sometimes with a sharp ridge marking the junction with 
the obconical tube. Fruit broadly obconical, 5-8 x 5-7 mm., the calyehie and 
staminal rings forming a Hal band or dark line around the outside of the rim; 
disc rounded, half to one nun. broad, valves short and broad. in orifice (pi. itt, 
fig. 3). 

Occurs in the Mount Lofty Range and south to Mount Gambier and lo Victor 
Harbour. Also on Kangaroo Island. 

17a. Eucalyptus ovata var. ukanuikloua Maiden in CVit. Rev., 27, 146 (1916). 

Ruds (15 x 9 mm.) and fruits (10 x 13 mm.) larger than above, the forma 
with rostrate operculum and the above mentioned ridge, the latter broadly 
obconical and slightly constricted below the rim. 

Kalangadoo and Mount Gambier. 

18. Eucalyptus elaeophora E. Muell in Fragm., Iv, 52 (1864). 

Medium to large-sized tree. Mature leaves alternate, petiolate, lanceolate to 
hroadly lanceolate, 10-22 x 1*5-3 cm. Umbels axillary, sometimes in pairs, 3-7- 
flowered, peduncles flattened and angular, 5-12 mm. long. Buds sessile or sub- 


sessile on thick angular base almost indistinguishable from the tube, cylindrical, 
more or less bicostate or angular, 8-10 x 4-6 mm. Operculum conical, shorter than 
lube. Fruit sessile or very shortly pedicellate, cylindrical or ohconical with a thick 
more or less angular base, otherwise smooth, 6-9 x 6-8 mm.;- disc less than 1 mm. 
broad, forming a narrow rim around the short broad valves which arc exsert or in 
the orifice {pi. iii, tig. 4). 

Blakeiy apparently had no South Australian material at Sydney except the 
poor specimen which was included under /;, tfoniocatyx. so reference to this State 
was omitted on p. 146, but il was included in rhe list of species occurring in South 
Australia on p. 289. 

Southern Flinders Range, Mount T.ofty R3ti£e and southern parts of the 
Slate, including Kangaroo Island. 

19. EuovLvrxus cosmofuvlla F. Mucll. in Trans. Viet. Inst., 1, 32 (1855). 

Varying from a small mallec-lilcc shrub to a medium-sized tree. Mature 
leaves alternate, petiolate. usually thick and coarse, lanceolate to broadly lanceo- 
late 10-15 x 1-4 cm, Umheh axillary, 1-3-llowered, peduncles very short and 
sometimes Inciting. Buds ubovoid. bicostate on very short flattened pedicels, 
14-18 x 9-12 nun. Operculum conical, conieul-apiuilate or rostrate, shorter than 
or equal to tube and sometimes wider. Filaments cream or pinkish coloured. 
fruit hemispherical, turbinate or shortly cylindrical, more or less bicostate, 
10-1/ x 12-18 mm., the calycine and stamina! rings forming a sloping outer edge 
to the rim, with the steeply oblique disc lining the inside around Hie 4-6 short, 
broad bur acute valves which arc just below the orifice (pi. iii, rig. 5). 

Blakeiy states thai the umbels arc hnl none with more than 
three flowers have been seen by the writer. Mueller's original description was 
"tri-Horis." Neither of* Muellers variations, included by Blakeiy (p. J 03-) are 
sufficiently distinct In warrant separation. The figures m Critical Revision 
(pb 91,. fig. 4-5) show only slight deviation from the normal, and recent collect- 
ing does not indicate any real differentiation. 

Occurs in the Mount Lofty "Range and south to Encounter Bay and Kan- 
garoo Island. 

20. Eucalyptus pniriNiANA Maiden in Crit. Rev. 16,211 (1912). 

Malice. Mature Idn-rs alternate, petiolate. lanceolate to broadly lanceolate, 
6-9 x 2-4 cm, Umbels axillnry, 3-8-flowcred, peduncles terete or scmiterete, 
clellcxed. B%$$ ohconical or cylindrical, rostrate, pedicellate, striate, 20-30 x 
10-14 mm. Operculum hemispherical-rostrate or conical rostrate, smooth or 
faintly striate, Fruit pedicellate, cylindrical and gradually narrowing tuto the 
pedicel, striate, disc as described below for E. incrassata, valves with short points, 
deeply enclosed (pi. iii. fig. 6). 

Occurs on the eastern border of the Nullarbor Plain at Ooldea and Barton. 

21. Eucalyptus uncuassata LabUt. in Nov., Holl., pi. ii, 12, t. 150 (1806), stricfo. 

Malice, Mature leaves alternate, petiolate, narrow to broadly lanceolate, 
6-12 x 1 *5-3 cm. Umbels axillary, 3-7-flowcred, peduncles slender., terxite or semi- 
tereie and slightly angular, up to 3 cm. long. Buds cylindrieal-obovoid to urceolate- 
rosiraie, shortly pedicellate in the former case to long pedicellate (up to 6 mm.) 
in the latter, 10-18 x 5-7 mm. Operculum conical to very rostrate with a more or 
less hooked apex, commonly somewhat wider than the calyx tube and shorter, 
smooth or with obscure fine wrinkles or with promiuent lines but not distinctly 
ribbed. Tube ohconical or constricted in. the middle and verv urccolate, smooth 
to striate with fine lines or obscure ribs. Fruit pedicellate or "sessile, urccolate or 
barrel-shaped being- constricted at each cud, smooth or slightly wrinkled M-hen 


mature, 10-12 x 9-10 mm.; the tube wider than the disc, so that the calycinc fcinfe 
forms a narrow but distinct platform around the erect rather sharp disc whose 
outer face is formed by the old staminal ring and whose inner is oblique to almost 
erect; valves with subulate points due to the persistence of the base of the style, 
when open either in orifice or, more usually, below ami enclosed (pi. ii, fig. 8-9 j. 
When immature the top of the ovary is ridged between the unopened valves. This 
last character is found in the remaining varieties below, and also in the member 
of the thtmosa complex. 

Occurs on Yorke Peninsula, in the Flinders Range, the Murray Malice and 
.south lo the Upper South-Kant 

It has proved inrposMuk: to make a satisfactory demarcation between litis 
species and E. angutosa Sehauer,. a wide range of intermediates having- been 
collected. The specimens have therefore been grouped as follows. It is probable 
that these represent mere points in a series, and collectors are warned that inter- 
mediates may be found. 

21a. Eccalvptus incrassata vaiv cost ata N. T. Burbidge stat. nov,: 
E. costata et. Muell. in Miq. Ned. Kruidk Archief., 4, 136 

A coarser form than the above. Mature hares fairly thick and coriaceous. 
Umbels on slender flattened peduncles (up to 3 cm. lung), which broaden below 
the flowers. Buds similar to the above, hut more definitely ribbed or striaie. 
Operculum as above. -Fruit on short, more or less angular pedicels, barrel- 
shaped or cylindrical, marked by sirialinns or sharp narrow ribs, 10-18 x 9-12 mm. ; 
disc as above, valves enclosed (p1. if, tig'. 10). 

This is the R. ancjulosa of various writers on South Australian ecology 
(10) (19). J. M. Black combines it with the following as did Maiden. The 
striation or ribbing of both the hud and the fruit is far less pronounced hi fresh 
material. The ribs are revealed on drying, due to shrinkage of the les? scleriscd 
portions. Maiden (pi, 14, fig-. 2 ) shows a fruit rather more coarsely ribbed fhaa 
the majority of specimens, hut it can still be contrasted with that in his fig. 5> 
which is the form described below. 

Occurs on Eyre and Yorke Peninsular and Kangaroo Island; also in the 
Murray Malice and the Upper South-East, 

21b. Eucalyptus ixc'uassata var. angulosa Beuth. in Fl. Aush, iii; 231 (1866), 
pro parte; E. ungttlosa Schauer. in Walp. Repert., ii, 925 (1843) sevsu 

A very course form with thick lanceolate leaves. Umbels 2-5 flowered, 
peduncles thick, flattened. 10-20 mm. iong. Buds obovii.e or cylindrical wilh 
short pedicels, 20-25 x 10-15 mm. Operculum conical or shortly rostrate, striate 
or ribbed, shorter hut not wider than tube. Tube ohconical or cylindrical, coarsely 
and deeply ribbed by projecting" hard ridges which continue into the slightly 
flattened and ribbsd pedicel. Fruit almost cupular to cylindrical, coarsely ribbed 
by the abovementioned ridges, 15-20 x 14-1S mm.; disc and valves as in other 
forms above (pi. ii, fig, 11). 

This coarse form agrees with Maiden's illustration (pi. 14. fig, 1, 3, and 54, 
i.e., with Western and South Australian material. Tn this Stale it is mainly found 
on Eyre Peninsula. 


This group of forms has always been a difficult one. The type of E, dumasa 
was collected by Allan Cunningham in New South Wales. Specimens in the 
Sydney Herbarium, collected at Wyalong. which is accepted as approximately the 
type locality, proved to have cylindrical buds which were smooth or almost so, 


find to have a conical operculum shortvr than the calyx tube. This form is rare 
i3i South Australia but occurs atpfljj the eastern border and near the Murray 
River. In this State most of the material classed as E. dunwsa has hurls with 
ribbed conical upereula. This agrees with the. Western Australian species, 
£. plleata. There arc imennctliates In the eastern part of the State, but on the 
whole the two are reasonably distinct. There is also E. brachyealyx with an 
obtuse, conical opcreuhmi which is striate rather lhan ribbed. On Kangaroo Inland 
there ;are a number of form*. The most distinctive is R canglobatu, in which the 
buds are so sessile as to be flatlcned at the base and the calyx tube is wider than 
long. There is also E. rugosa, where the bud has a flattened pedicel and the. tube 
is 2-4-ribbed, 

Unfortunately, the whole group is complicated by the description of 5, anceps. 
Maiden described this as a variety p{ E. conc/toba'tci, bin placed it inOMm-.-dia.te 
between fr conglobota and E. duniosa. It differs from the former in that the 
buds and [rQtfo though sessile, are not hasally flattened and the fruits are barrel- 
shaped and from E. dtimosa in a relatively shorter calyx tube and in the 
lack of pedicels. Compared with true R. dumosa this diagnosis would hold, but 
in practice E. weeps lies between E. pileabt, II. rutjosa and E. conglobata, and as 
such it includes a heterogeneous collection of variaiions, Presence or absence of 
pedicels Is not a good diagnostic, character, and sessile buds and fruits oE E. rituosa 
are difficult to place; the .same applies to E. plfeuta, and apart from individualistic 
variations such as these there are probably a number of hybrids and eeoiype^ 

Auother species related to the group but more easily recognised is E. coiia'nna. 
An effort to reduce .same ot" the forms to varieties proved unsatisfactory and 
they were retained a.s species to avoid adding to the existing confusion. 

22. Ft'CALYPTOs conglobata (R. Br.) Maiden in Grit. Rev., 6, 273 (1022). 

Ma-lee, Mahtrc leaves alternate, peUolalc, thick, narrow to broad lanceolate, 
5-14 x I -5-2-5 cm. Umbels axillary, 3-8-fiowcrcd, peduncles short, thick, more 
or less angular, broadened at the top to form a receptacle, Buds squat, broadly 
ovoid, flattened at the base and closely sessile, 8-10 x 6-8 mm. Operculum shortly 
and broadly conical, obtuse or acute, smooth or slightly striate, as long as or 
.slightly longer than tube. Tube shortly cylindrical, bicostate, about twice as wide 
as lone;. Emit cylindrical or hemispherical, 2-3-ribbed, 5-6 x 8-10 mm., disc 
narrow, valves with subulale points which project above the rim (pi. lit, Jl*. 7). 

Eyre Peninsula and Kangaroo Island, 

2,1 Eucalyptus Aisrcr.i's (R. Br.) filakcly in Key to Su& a US (1934), 

Malice. Juvenile leaves opposite, sessile, slightly lobed at base as to be 
almost stem clasping, ovale to broadly lanceolate. Mature lanvs alternate, 
pctiolate, lanceolate, rather thick and rigid. 5-12 x 1-2*5 cm. Umbels axillary,, 
3-6-flowered, peduncles relatively thick, flattened or subareolar, 6-12 mm. long. 
Htuis tiesj&t or almost so, cylindrical or ovoid, narrowed at both ends, 8 ID x 
4-6 mm. Opereulum conical, obtuse. acute or shortly rostrate, radialely striate, 
about the same length as the lube or slightly shoitcr. Fruit sessile on thickened 
peduncles, cylindrical to barrel shaped, striate, 7-10 x 6-8 mm, : jifcc narrow, valves 
in orifice, short but subulate (pi. iii. fig. 8). 

Eyre Peninsula, Ynrke Peninsula, Murray Ma!lee, Upper Soulh-Easl and 
Kangaroo Island. 

24. Eucalyptus urtios a (R. Br.) Hlakely in Key to TXuc. 120 t.1934). 

MaUec. Mature le>wes narrow to broadly lanceolate, 5-10 x 1-2 cm. Umbels 
axillary, o-8-flowered, peduncles stout, flattened ^ broadened below flower^ becom- 
ing coarser during itmlnratiun oi' fruit.--. P>ods pyriform on short, stout, ilatiened. 


3-4-augled pedicels* 6-7 (9-12 with pedicels) x 5-6 mm, Operculum hemispherical, 
flat-topped or very obtuse, smooth or slightly striate, a little narrower than the 
ribbed tube or almost equal, also shorter, fruit pyriiorm or turbinate, ribbed -with 
2-4 main ribs two of which are broadened below into the flat pedicel, and some- 
limes with a lew striatious (pi. iu\ fig. 10). 

Eyre Peninsula, Kangaroo Island, Encounter Bay and the Upper South-East. 

25. Eucalyptus jhjmos* A. Gain, fide Schauer hi Walp, Rep. Hot, Sy*t., 2, 925 


Malice. Mature leaves alternate, pefiolate r lanceolate, 6-10 x 1-2 em. Umbels 
axillary, 4-S-flowered, peduncles- comparatively stent and scmitereic. Buds 
smouth, cylindrical, shortly pedicellate. 7-12 x 3-5 mm. Operculum hemispherical 
to shortly conical, minutely striate or almost smooth, much shorter than the. 
cvindrical tube. Fruit cylindrical to eampanulate, somewhat wrinkled or striate, 
.shortly pedicellate, 6-10 x 5-7 mm.; disc narrow, valves with protruding subulate 
points (pi. in, Eg, 9). 

A New South Wales species which extends to the eastern border of this Suite 
and down into the Murray Mallee. 

26. Eucalyptus pileata Btakcty in Key to Kuc )( 120 (1934). 

Mallee, Mature leaves alternate, petiolate, narrow to broad lanceolate. 
5*5-10 x 1-3 cm. Umbels axillary, 3-f>flowered, peduncles terete or semiterete, 
6-15 mitt. long. Bttrfs pedicellate, obovoid or cylindrical, 8-14 x 4-5 mm. 
Opereulum hemispherical to acutely conical, radiately striate or ribbed, sometimes 
wider than tube, shorter than or equal to the cylindrical or obconical or cupular 
lube, which is smooth or striate. Fruit pedicellate, cylindrical or obconical. striate 
or almost smooth, 7-10 x 6-S mm. ; disc narrow,, valves sunken with protruding 
subulate points (pi. hi, hg. 11). 

Eyre Peninsula. Yorke Peninsula, Murray Mallee, Encounter Bay, and Kan- 
garoo Island. 

Form a. Obovoid pedicellate buds with almost smooth operculum. No Droits 
available. This form seems distinct enough to warrant mention, but without 
fruits and further information it docs not seem strong enough to make a variety. 
It occurs on Eyre Peninsula. 

27. Eucalyptus brachycalyx Rlakely iu Key to Euc, t 119 (1934). 

Mallee. Mature leaves alternate, petiolate, linear to narrow lanceolate* 
shining. .5-10 x 0'7-l*5 cin. Umbels axillary, 3-9-flowered. peduncles slender, 
slightly flattened. Buds ovoid-ellipsoidal, ou slender pedicels* 7-9 x 4-5 mm. 
Open ttiuiU conical, obtuse, about the same size and shape as the cupular, striate 
tube, usually with irregular wrinkles rather than striate, commonly slightly 
different in colour from the tube. Fruil hemispherical to eampanulate or turbinate, 
smooth or faintly striate, 6-7 x 5-6 mm.; disc narrow, less than 0*5 mm. wide; 
valves subulate and exsert except where the brittle points have been broken oft" 
tc.f. Blakely) (pi, iii, fig. 12 J. 

This species could be mistaken for JE, oleosa var. angiu>l-ifaHa } but it can be 
differentiated by the large obovate versatile anthers with a conspicuous gland and, 
in fruit, by the turbinate open capsule (not globular as in E. oleosa) and by the 
spreading rather than erect valves. 

Eyre Peninsula, Flinders Range and to the east thereof aV>ng I he border of 
the shrub steppe and mallee scrub. 

27a. Elxalyptus brachycalyx var. chinuoo Blakely l.c. 

Buds somewhat smaller than those above, 5-6 x 3-4 mm. Operculum striate 
and very obtuse, slightly longer rhan substriate cupular lube, also slightly narrower 
than tube giving a marked "egg-in-egg-cup" appearance. Fruit as above. 


It is doubtful whether this variety is a true one. The species itself is quite 
distinct and may be recognised by the slender peduncles and pedicels lo the uwheU, 
the cupular calyx tube about the same size and shape as the operculum and by 
the narrow shining leaves. It was unfortunate that Blakely contradicted his own 
description of the operculum. 

Distribution as above. 

28. Eucalyptus cokcinna Maiden and Blakelv in Crit. Rev., 71, 49 (1929) ; 

E. ochrophylla Maiden and Blakely in Crit. Rev., 71. 50 (1929). 

Malice, Mature leaves alternate, petiolate, lanceolate, broad or narrow, thick 
and glotsy. Umbels axillary. 3-7-flowered. Peduncles terete or semi terete, 
6-12 mm. long:. Buds clavate, pyriform or cylindrical, 8-10 x fi-K mm. on pedicels 
like the peduncles and 4-12 mm. long. Operculum hemispherical, very obtuse or 
flat-topped, about half as long as broad, more or less striate, sometimes slightly 
broader than tube. Stamens red (type from Victoria Desert) or pale (Cleland's 
specimens quoted under original description ot E. ochrophylla) . Fruit pyriform, 
almost turbinate or cylindrical, smooth or faintly striate, 6-10 x 6-10 mm.; disc 
about 1 mm. wide, rounded above with the dark line of the calycine ring outside, 
capsule slightly sunken but the subulate valves protruding and spreading' or rarely 
coherent at top as in E. oleosa (pi. iii, fig. 13). 

Clcland's specimens of E. ochrophylla show a definite variation towards 
E, concinna, and the buds cannot be separated in any particular character. The 
type of E. concinna has very long pedicels (matched among E. ochrophylla 
material) and the buds are golden-brown, whereas in E. ochrophylla they are 
greenish. The yellowish colour of the leaves in the latter species is not q reliable 

Occurs along the transcontinental railway al Ooldca and Innnana. 

29. Eucalyptus T.KPTOPitvu.A F. Muell. in Miq, Ned. Kruidk. Archief., 4. 123 


Malice, Mature leaves alternate, petiolate, linear or narrow lanceolate, with 
numerous oil glands., 5-10 x 0"6-l'5 cm. Umbels axillary, 3-8-flowered peduncles 
5-8 mm. long, terete or semitercte. Buds like two more or less equal colics 
attached base to base, on a slightly angular pedicel as thick as the peduncle, 
5-7 x 3-4 rnm J Operculum conical* acute, or obtuse, about as long as tube or 
-ihorter. Stamens sharply reflexcd in bud and with a sharp hend in the filament 
when (lowering, fruit pedicellate, hemispherical (o cupular, 4-5 x 4-5 mm,; disc 
forming a flat or rounded ring at the Lop of (he fruit ; valves narrow, subulate and 
protruding through the orifice (broken off in old fruits) (pi. iv, rig. 12). 

Along - the transcontinental railway at Ooldca and Tareoola, also Eyre and 
Yorkc Peninsulas. Flinders Range, Murray Malice and Kangaroo Island. 

.10. Eucalyptus uncixAta Tun:*. Bull- Snc Nat. Mnpc, 22, 23 (1849). 

Very close to the preceding species but buds and fruits somewhat coarser. 
The chief di (Terence is in the bud, which is sessile or on a short thick pedicel 
indistinguishable from the tube. Operculum conical, obtuse and shorter than the 
obconieal or cylindrical tube 6-8 x 3-4 mm. fruit sessile or almost so, pyriform 
or barrel-shaped, 5-6 x 4-5 nun. (pi. iv, fig\ 13). 

These two species were united under the latter name hy Benthatn in the 
Flora Australiensis, but were separated by Maideit. It would be difficult to 
separate them without buds, though the slender pedicels and ycllow-rcd hranehlets 
in E r leptophylla ate characteristic. 

Eyre Peninsula seems to mark the eastern limit of this Western Australian 



31. Eucalyptus cneorifolia DC, Prodr,, 3, 220 (1828). 

Mallec. Mature leaves alternate, petiolate, narrow linear to linear lanceolate, 
5-11 x 0'4-l cm. Umbels axillary, 6-1 5-fiowcrcd , peduncles short. Buds cylin- 
drical, shortly pedicellate or sessile, 7-12 x 3-5 mm. Operculum smooth, cylin- 
drical to conical, obtuse, about the same length and shape as the tube. Frail 
shortly pedicellate or sessile in densely packed swni globular clusters, hemispherical, 
5-6 x 7-8 mm.; disc flat, about 1 mm. broad, around the partially exsert, subulate 
but short valves. 

Kangaroo Island and Encounter Bay. 

32. Eucalyptus Gillii Maiden in Crir. Rev., 15, 177 (1912). 

Glaucous malice. Mature leaves opposite or almost so, glaucous, sessile Or 
shortly pedicellate, ovate, oblong or shortly lanceolate, 4-6 x 1-2-5 cm. Umbels 
axillary, glaucous, 4-9-flowered, peduncles scmitcrcte, slender. Buds glaucous, 
ovoid or cylindrical-rostrate, pedicellate, 8-15 x 4-7 mm. Operculum conical, 
acuminate or rostrate, longer than trie cylindrical tube, fndl ovoid or globular, 
pedicellate, 5-8 x 5-8 mm.; disc very small, valves exsert. long and subulate but 
usually broken oft" short. 

Northern Flinders Ranges. 

32a. Eucalyptus Gillk var, petjolaris Maiden in Jour. Roy Soc. N.S.W., 
53,59 (1919). 

Differs from the above only in that the leaves are definitely petiolatc (up to 
1 cm.) and broadly lanceolate. It is doubtful whether this form is worth retaining 
as a variety. 

The type came from Wirrahara, but other specimens from the northern 
Flinders Range have been seen. 

33. Eucalyptus oleosa F. Muell. in Miq. Ned. Kntidk. Archief,, 4, 128 (1856), 

sensu stricto. 

Malice or small tree. Mature leaves alternate, petiolate, narrow lanceolate, 
usually glossy, 5-10 x 1-2 cm. Umbels axillary, 5-14-flowered, peduncles semi- 
terete. Buds pedicellate, ovoid or cylindrical. Operculum conical, longer than 
cupular or semiglobular tube. Fruit pedicellate, globular or clavate 5-9 x 5-9 mm.; 
disc narrow, valves thin, long and subulate, very exsert. coherent at the apex but 
breaking easily and hence often lacking from specimens (pi. iv, Cg. 4). 

A photograph and some fragments of the type material have been obtained 
through the courtesy of Dr. F. P, Junker of the herbarium of the Stave University 
at Utrecht, Holland. It is mixed E. uncinata Turcz. (buds and flowers) and 
E. oleosa F. Muell (immature fruits). The latter, which is regarded as the true 
type, was, according to the Melbourne Herbarium authorities, collected from the 
Murray Malice, When inquiries concerning the type were first made it was hoped 
that the specimen would show the shape of tlifc operculum, since this feature is 
variable and has been the basis of differentiation in several varieties. Unfor- 
tunately, this is still unsettled, it is believed, however, that the specific name 
was associated with a form having a conical operculum, since Mueller's species, 
E\ sOci<t!is t published in the same paper, has a rostrate one. 

< Jne of the most important and widespread mallee species, as can be seen by 
reference to Wood's vegetation map (10). 

33a. Eucalyptus oleosa var. Peenetu Blakely in Key to Euc, 270 (1934). 

Leaves, branchlets, buds and fruits more or less ashy -grey. Leaves ob'onq 
to oblong lanceolate. Operculum obtusely conical, twice as long as cupular tube. 

This may not be a good variety. Blakely founded it on material from Oohlea, 
and specimens probably belonging to it have been received front F.yre Peninsula 
and Kangaroo Island, 


33b. Eucalyptus olkosa var. gf.auca Maiden in Jour. W. Aust. Nat, Hist. Soc. t 
3 t 171 (1911); E. socialis F. Mucll. in Miq. Ned. Kruidk. Archief., 4, 
132 (1856); £. trcwscontincntalis Maiden in Jour, Roy. Soc. N.5.W-, 
53,58 (1919). 

Leavfis lanceolate to broad lanceolate, usually dull or subgtaucous- Buds 
cylindrical* or urccolate-rostraie Operculum rostrate and longer than cylindrical 
or urceolate tube. Fruit narrowed into orifice and sometimes with a tendency 
\n be urceolate (pi. iv, fxpf- 1 ) - , 

The nomenclature of this form was confused by Maiden. In Cnt. Rev., 15, 
167, he redescribed var. gtauca and listed E. sodaHs F, Muell. as a synonym. 
Later he quotes his description of E, trancontinentalis (Crit. Rev., 34, 268)wttb 
E. oleosa var. glauca as a synonym, but ihcrc is no mention o£ E. sociahs. South 
Australian material matched with Maiden's specimens in Sydney is identical with 
the type of E. socialis from Mueller 1 * material in the Melbourne herbarium. It 
does not seem to be distinct enough for specific rank and forms a link between 
E. oleosa and E. Flocktoniae. It would appear that Blakely's description of 
11. socialis was superfluous. 

This form occurs from the Nullarbor Plains eastwards across Eyre and 
Yorke Peninsulas, along the Flinders Range to the north and also east to the 
Murray Mallee, the Victorian border and Kangaroo Island. 
33c EucALYrTUS oleosa var. angustifolia Maiden in Crit, Rev., 39, 278 

Leaves narrow to broad lanceolate, usually glossy. Buds pedicellate, cylin- 
drical or ovate, Operculum crumpled or wrmlded on the surface, conical or 
cylindrical, very obtuse or flat-Lopped, different in colour from the tube, Fruit 
globular (pi. iv, (ig. 2"). ( 

When Maiden described this variety he gave £. socialis F. Muell. as a 
synouvm and figured (Crit. Rev., pi. 65, tig. 17 a-b) a Murray River specimen 
collected by Mueller. This specimen is referred io as '"the type,'* and then as 
"a repuWtype specimen from Mueller," but it is not cleat whether he means it 
as a type of 'E. socialis or of his var. ancjustifolia. There is, however, no mention 
of such a specimen under the original description of £. socialis, so that E. socialis 
is not a svnonym of this variety. Blakely gave Mueller's species full ranking, but 
retained E. oleosa var. annuslifolia and quoted a specimen from Pinnaroo, collected 
by J. M. Black, as the type, 

Subsequent to the publication of the Key, Hlakcly appears to have decided 
that E. oleosa var. cwgusiifolia should be given specific rank. There is a large 
umount of material in the Sydney Herbarium which he separated out under the 
name E. lamptophylla. This includes Black's Pinna ron specimen. The species 
was never published, it is a distinct variety but, without the characteristic buds, 
difficult to separate from the rest of £. oleosa. Since there is doubt concerning: 
the bud shape in the original E. oleosa, specific ranking for this form would 
hardly be advisable. Mueller's specimen* mentioned by Maiden, is presumably 
the type. 

Distribution is the same as for E. oleosa sens-it stricto. It is particularly 
common on l£yre Peninsula and Kangaroo Island. 

34 Fuc^Lvrrus Fuicktoxiae Maiden in Tour. Roy. Soc. N.S.W., 49, 316 

Malice or small tree. Mature leaves alternate, petiolatc, lanceolate, 6-12 x 
1 -5-2'5 cm Umbels axillary, 3-7-flowered, peduncles semiterete. broadened at 
top "but not flattened. Buds urceolate- rostrate, pedicellate, 12-17 x 5-8 mm. 
Operculum with broad base, hemispherical or conical and with a totig beak, 


up to twice as long as the urceolatc tube and broader at the junction. 
Fruit urceolatc, broad at base and constricted below the wide orifice; disc narrow, 
valves as in Bi oleosa but very brittie and usually broken off and hence appearing 
enclosed, 8-10 x 7-9 mm. ; pedicels angular (pi. iv, fig. 3), 

This species forms a link between £. oleosa and the Western Australian 
species &] torquata. It occurs from the Nullarbnr Plain across Eyre and Yorke 

35. Eucalyptus calycocona Turcz. in Bull. Pbvs. Math. Acad. Petersb., 10, 

338 (1852). 

Malice. Mature leaves alternate, petiolate, linear to lanceolate, 5-10 x 
0*7-2 cm. Umbels axillary, 3-7-flowered, peduncles slender, terete or semiterete. 
Buds clavate, quadrangular, pedicellate, 6-10 x 4-6 mm.; with numerous oil 
glands. Operculum shortly conical, obtuse, acute or apiculate, not angular, shorter 
than the angular tube. Fruit cylindrical or urceolate, quadrangular but less 
sharply so than when in flower, smooth, pedicellate, 8-12 x 5-6 mm.: disc lining 
the top of the tube and obscured by the narrow persistent staminal ring; valves 
small and deeply enclosed (pi. iii, tig. 15). 

Eyre and Yorke Peninsula, Murray Mallec and upper South-East, 

35a. Eucalyptus calycogona var. Stakkortmi Hlakclv in Key to Euc., 265 

A coarser form than the above. Mature leaves lanceolate to broadly 
lanceolate, thick, 6-12 x 1-5-2*5 cm. Umbels as above. Buds 15 x 5-6 mm!, 
including the pedicel, shape as above. Operculum conical acute, apex angular. 
Stamens white or pink. Fruit 14-16 x 7-9 mm, 

Eyre Peninsula and, according to Blakely, also near Encounter Bay. 

36. Eucalyptus gracilis F. Muell. in Trans. Vict Inst, 1, 35 (1855). 
Mallee or medium-sized tree, Mature leaves alternate, petiolate, narrow 

linear to lanceolate, oil glands conspicuous as black dots, 4-9 x 0*6-1 '5 cm. 
Umbels axillary. 4-12-flowered, peduncles terete or semiterete. Buds clavate 
(with pedicel), not angular, with numerous oil glands, pedicellate, 3-5 x 4-5 mm. 
(pedicel about 3 mm). Operculum hemispherical or patelliform, obtuse or 
apiculate, shorter than the obconical tube. Fruit ovoid to pyriform, smooth, 
pedicellate, 5-6 x 3-5 mm.; disc lining the top of the tube or oblique, the rim 
formed by the narrow ridge of the stamina) ring; valves short and broad, enclosed 
or very near the orifice (pi. iii, fig. 14). 

Eyre and Yorke Peninsulas, Flinders Range and south to the Clare hills, 
Murray Atallee and upper South-East. 

36a. Eucalyptus gracilis var r ekkcta Blakely in Key to Euc, 266 (1934). 

Loaves erect, narrow linear, shining, oil glands very numerous. This appears 
to be a dry country form. It is possible that it is not a very definite variety. 

Eyre Peninsula, Murray MatJee and the northern edge of the mallec scrub. 

37. Eucalyptus intertexta R t T. Baker in Proc Linn. Soc. N.SAW, 25, 308 

Small to fairly large tree. Mahtre leaves alternate, petiolate, linear to lanceo- 
late, 7-12 x 0'7-2 cm., bluish or stibglaucous. Umbels in terminal or subterminal 
panicles or rarely solitary iu upper axils, 4-6-flowered, peduncles slender, terete 
or semiterete. Ends ohovofd, clavate on slender, subangular pedicels, 5-7 x 
3-5 mm. Operculum conical, acute, somewhat narrower than the tube and shorter. 
Tube obconical and sometimes slightly constricted in to the orifice, with one or 
two lines continuous with the angles on the pedicels. Frmt obovoid or pvriforrn, 


pedicellate 6-8 x 5-7 mm.; orifice contracted, (he persistent statmnal ring com- 
moulv hiding the narrow disc; valves short, enclosed or near (pi iv, hg. 14). 

Everard and Birksgate Ranges and south-eastwards to the Hinders Range, 
also west of Port Augusta. 

37a Eucalyptus jntektrxta var. fruticosa Blakely and Jacobs in Key to 
Euc. 168 (1934). 

Mallee Mature leaves as above but rarely dull and hluish. more usual y 
shining and vellowtsh-green when dried. Umbels as above Operculum shortly 
conical and sometimes apiculate, due to the point on the thin pre-opercuHuil which 
persists till or almost until flowering. Fruit as above. 

This form is found on rock slopes in arid country, whereas the typical form 
occurs low on the slopes adjacent to creeks bordered by E, camaldnlensis. It was 
based on a Northern Territory specimen but extends down to the northern 
Flinders Range around Hawker and Quom. 
38. Eucalyptus coot.abah Blakely and Jacobs in Key to Euc, 245 (1934). 

Tree Mature leaves alternate, petiolatc, narrow to broad laneolate, 7-15 x 
1-3 cm. Umbels in short terminal or subterm'mal panicles, 3-6-flowercd, 
peduncles slender, subangular. Buds ovoid, pedicellate, 3-5 x 3-4 mm Operculum 
conical, acute or apiculate. about the same length as tube or slightly longer i-rn/f 
hemispherical or broadly turbinate, 3-4 x 4-5 mm., the rim of the tube th.n and 
fragile with very thin disc; valves short, broad, obtuse, exsert and more or less 

incurve^ ^^ Australian mate rial of ihis species was formerly placed with 
E microtiieca F. Muell. The differences between the two appear to be slight. 
It' occurs in the northern parts of the State near I-akc Eyre and the northern 
Flinders Range. 

39. Eucalyptus letjcoxylon F. Muell. in Trans. Vict. Inst., 1, 33 (lUbb), 
sensu stricto. 
Smalt or medium-sized tree. Mature leaves alternate, petiolate, narrow to 
broad lanceolate, 7-15 x 1-3 cm. Umbels axillary or lateral on leafless portions of 
branchlets or in short panicles. 3-floweted (except variety), peduncles terete 
or semitcrete. Buds clavate, ovoid or cylindrical-rostrate, wrinkled when dry, 
pedicels normally long and flattened and angular above as they pass into the calyx 
tube sometimes' short and semilercte. Operculum conical, acute or rostrate with 
a sometimes angular point, as long as or longer than the obcomcal or turbinate 
tube Stamens with translucent oil glands in the filaments, especially the outer 
ananthe.rous ones, Stamtual ring 1-2 mm. wide, conspicuous when the stamens 
fall, hiding the oblique disc, deciduous during the maturation of the _ fruit or 
remaining attached to one sale of the nm. Fruit pyriform, ovoid or suhglobuar 
on Ion" terete or semitcrete pedicels (except in short pedicetled forms); disc 
oblique'but not covering the short broad enclosed valves. In (he typical lorm he 
buds are 8-13 * 6-7 mm. and the fruits 10-15 x 8-13 mm, (pi u, ng. 7). Tins 
occurs in the southern Flinders Range, Mount Lotty Range, Murray lands and 
in the South-East and on Kangaroo Island. 

This species shows considerable variation and a large number of t otitis have 
been named Both red- and white-flowered individuals occur, but are not 
restricted Lo any particular variety. Some plants are glaucous, hut not reliably so. 
Many of the forms seem to grade into one another, hut despite this the following 
varieties are included. 
39a. Eucalyptus i.mrcoXYi ois var. mactiocaupa J. F., Brown m For. H, 

S Aust (1882; £. f.i<u.nx\<lo>i var. erylhronenm F, Much, in Mkj. Ned 

knndk. Archief,. 4, 127 (1856). 


Buas 16-20 x 10 mm., pedicels very angular. Operculum rostrate. .Fruit, 
15-22 x 15-18 mm. (pi. ii, fig, 6). 

There seems to be some room for doubt concerning the legitimacy of Brown's 
name. Maiden (Grit. Rev., 21, 91) accepts it because the variety includes both 
red- an-! white-flowered terms. This point had been mentioned in Brown's 
description, where it is stated that "we have ventured to applv the name macr»- 
carpa .... as being- more directly specific; and, in order to individuate them from 
the red-flowering sports: of the true representative of the species Lcucoxyion." 
Thisiatterfact is the more important as it means that Mueller's description is 
definitely inadequate, being "filamentis sanguineis." According to Maiden 
(I.e.) and Brown there is no doubt as ro the synonymity of the two names, 
Maiden and Elakely have been followed in this paper because: (i) Mueller's 
description is inadequate; (ii) the -wide variation within the species means that 
varietal names represent little more than arbitary points in a series; and (iii) so 
little can be gained in this case by a strict application of the rules of priorilv. 
To waive Mueller's description because it is not diagnostic is an admittedly risky 
procedure, since what constitutes an adequate description has never been definitely 
siatcd and the great majority of the early descriptions leave much to be desired. 
However, in this case an argument can be made for either name, and therefore the 
status quo has been maintained 

The variety was based on specimens from Eyre Peninsula. It also occurs in 
the Mount I.ofry Range and cm Kangaroo Island. Intermediates occur in the 
Flinders and Mount Lofty Ranges. 

39b. Eucalyptus letcoxvlon var. ancitlata Deuth. in R Austral., 3. 210 
( 1866) . 

The relation between this variety, the preceding and var. rugulosu F Muell 
is nijt dear (c.f. frit. Rev., 12, 91). Bent-ham's description states that the catyx 
tube, is distinctly angled. In most large-fluwered forms the pedicel is flattened 
as it passes into the tube and the two ribs thus formed, with two smaller one* 
between, pass upwards to be lust in the upper part of the tube. Blakely states 
that the fruits in this variety, which is not known to the writer, arc subangular. 
In all material available the fruits are rnunded. any angularitv having been 
during maturation. Maiden considered (I.e.) var. angulahi was not the same as 
vzr.macrocarpa, but gives no grounds for his opinion. Further collecting mav 
elucidate this point, but the variety does not seem to be a sound one. It is reported 
to have come from the Mount Eofty Range. 

39c. Eucalyptus LEucnxvT.oN var. PAdPEiurA J. E. Brown in For. Fl. S. Aust. 
(1882) ; E. jugalis Naudin in Second Mem., 37 (1891). 

Umbels three or more Qowered. Bliffa ovoid or pyriforin.. peduncles, and 
pedicels shorter than in above varieties, 5-7 x 3-5 nun. Operculum conical or 
comcal-apiculate rather (ban rostrate. Fruit globular turbinate to hemispherical, 
6-7 x 6-8 mm., relatively wider and shallower than in other varieties (pi ii, 
fig. 4-5). 

This form docs not seem to he sufficiently distinct to warrant specific rank. 
Some specimens are distinctive with more than three flowers per umbel and short 
pedieelled pyriform buds, but these grade into normal but small fruited forms of 
£\ leucoxylon. 

Mueller's original description of E. leucoxyhn stated "tri-raro-qumqtte- 
rlonV r This referred to his variety pluri flora which, as stated by Maiden, is 
E. caldculirix F. Muell, 

From Clare north into th* Flinders Range, also in the Mount Lofty Range 
and ou Eyre Peninsula. 

40 EucALvrrrs cAlcicuxtsix F. Mudl. ft Miq. Ned. Kruidk. Atchief., 4, 129 

Malice or small tree. Malum leaves alternate, petiohtc, lanceolate to broad 
lanceolate, the marginal nerve usually distant. Umbels axillary or clustered on 
leafless portions of the branchlets,3-7-fIowered, peduncles, scmiterete. 4-fc mm. 
tone 1 Puds davate on more or less angular pedicels 5-7 x 4-5 mm-. Ihe. surtacc 
crumpled or wrinkled like that of a withered and shrunken apple. Operculum 
conical, acute or obtuse, .lightly shorter than the tube Fruit pedicellate, some- 
times shortly so, pvriform or truncate-obovoid, 5-8 x 5-& mm.: disc narrow, very 
oblique or lining the top of LheJ.ube, valves short und enclosed but not so deeply 
so as in !S. odorata (pi. i, fig. 7), 

E cakkultrix occupies a position between £. odorata and die small fruited 
forms of Edeucox^lon, and it is significant that, while based nn Mueller s 
E. odorata var. cakkultrix, it also includes Ids E. kucoxylon var. pluriHora. %te 
species is a well-marked Otre and can be distinguished by the characteristic 
wrinkling or crumpling of the surface of the buds when dry, the usual retention 
of a stamina! ring during the maturation of the. fruit and by the marginal nerve 
of the lanceolate Teaf bem^ distant from the edge. HIakely's var. porosa based on 
E. porosa F. Mwell. illustrates the tendency towards E. kneoxyton and does not 
warrant separation. Var, obsatra appears to have been well named. There are 
two specimens in the Sydney Herbarium named R calacultrtx var. obsatra. Both 
were collected by Maiden at Eundaleer Forest, March 1397. One specimen is 
E fmlicetorum' 'which as stated below is E odorata var. angushfoUa, and this is 
also represented in the Melbourne Herbarium. The other appears to be that 
referred to as having "the general fades of E. fasckulosa" (Key to Flue.,. 2Zd). 
This latter specimen" probablv belongs with U. hcm'iphloia var. avcrofarpa. 

Eyre Peninsula, Yorke Peninsula, Flinders Kange, Adelaide .Flams, and 

Murray Mallee. 

The odorata comvlkx 

This, like Oft dnmasa complex,, has been the centre of .considerable confusion 
and the arrangement in the Kev to the Eucalypfs is unfortunate, The true situa- 
tion appears to be as follows: E. odorata, for which E, cajupttka I, Muell, is a 
synnnvm, is the tree form which is found around Adelaide and winch becomes 
malice-like as it goes north to Mount Remarkable and south to Encounter Bay. 
£. odorata var. anguslifolia is synonymous with E. frntkvtorum F. Muell, which 
is not distinct enough to warrant specific Tanking. U appears to link E. odorata 
with the very narrow-leaved fi. viridis R. T. Baker. The type specimen o.t 
Mueller's E, odorata var. erythrandra lias been seen. As poinrcd out. in a private 
communication by a Melhoume authority, it is the same as E. Lansdo-wneana 
F. Muell. and J. E. Brown. The latlci is a distinct species, having normally sessile 
and coarser hud* and fruits in short panicles or clustered umbels. s 
description of this variety is erroneous and appears to have been based on a red- 
flowered specimen of E. odorata. Tn the type specimens of K odorata (from 
Light's Pass) and of E, cafupuka (from the Flinders Kaujje) the filaments appear 
to have been red, though it is not always easy to be certain of this character when 
dealing with dried material. These specimens are in the Melbourne Herbarium 
and have been seen. Blakely's ft odorata var. rvfracia (Key to Euc 226) is an 
abnormality. Similar rerlexed stamens tipyt been observed in flowers of other 
species. Usually only some of the flowers on a branch are affected. This Variety 
cannot be retained. Var. macroenrpa (I.e.) is E. f.ansdomncana. 
4t. Kucalvttus Lansdownkana F, Muell. and T. E. Btown in For. Fl, S. Aust, 
pt be (1890). E odorata var. crvthrwidra F. Muelh in Mkj, Nod. Kruidk. 
Arch.rf ,4. 120 (1856). 


Malice. Mature leaves alternate, petioiate, thicker and coarser than those oE 
E. odorata, lanceolate to broad lanceolate, 8-13 x 1-35 cm. Umbels always 
clustered and usually in short panicles or the umbels grouped close together on a 
leafless portion of the stem, 3-8-flowcred, peduncles semheretc. Buds sessile oi on 
short thick angular pedicels diffiailt to distinguish from the obconical and some- 
what angular tube. Operculum conical acute or obtuse, shorter than tube. Fruit 
sessile or shortly pedicellate, cylindrical or barrel-shaped, striate or bicostute, 
8-10 x 7-9 mm.; disc very narrow, lining the top of the tube, valves short deeplv 
enclosed (pi. iv, fig. 8). : 

Southern Eyre Peninsula and Kangaroo Island. 

42. Eucalyptus odorata Behr and 5chlccht. in Linnaea, 20, 547 and 567 

(1847), sensn ftrkto; E. cajuputea F. Mudl. in Miq. Ned Kruidk 
Archte ., 4 f 126 (1856). 

Tree or mallee. Mature leaves alternate, petioiate, narrow to broad lanceolate 
(narrow m malice forms), 6-14 x 1-2*5 cm. Umbels axillary or clustered on 
leaner portions of the branchlets, peduncles semitcrctc, 8-15 mm. long. Buds on 
angular pedicels 2-7 mm. long and passing without abrupt change into rhe slightly 
angular obconical tube, 6-8 x 3-5 mm.; disc lining the ton of the tube, the rim 
thin, valves deeply enclosed, short (pi iv. fig. 5). 

Eyre and Yorke Peninsulas, southern Flinders Range, Adelaide Plains. 
Mount Lofty Range and south to Encounter Bay, Kangaroo Island and Murray 
Mallee. J 

42a Eucalyptus odorata van ancustifolia BJakely in Kev to Euc. 226 ( 1934) ; 
E. fruticetorum F, Muell. in Miq, Ned. Kruidk. Archief., 4. 131 (1856)! 
Mallee. Mature leaves alternate, petioiate, narrow lanceolate. 5-14 x 
Do-I-5 cm., surface dull and subglaucous, especially when young though older 
leaves are like of the above. Buds shortly pedicellate, smaller than in the 
above but subglaucous though similar in shape, fruit turbinate, cupular or 
pynform with striations running down into the short angular pedicel (pi iv 
hg. 6). ^ ' 

Port Lincoln, central Flinders Range between Quom and Wirrabara. 

43. Eucalyptus viainis R. T. Baker, in Proc. Linn. JJoc* N.S.W., 25, 316 (1900). 
Mallee or small poorly-shaped tree. Mature leaves alternate, petioiate, very 

narrow, linear, 6-16 x O3-0'5 cm. Umbels axillary, 3-8-flowered, peduncles 
setnitercte, 5-7 mm. long. Buds cylindrical or ovate", on short pedicels, 5-7 x 
3-5 mm. Operculum shortly conical, smooth or slightly striate but not pinched in 
to form an angular point, as long as the tube which is cylindrical and which pusses 
into the pedicel more abruptly than in any other members of the odorata complex. 
Frwt pedicellate, hemispherical to semi-globular,. 4-G x 4-5 mm.; disc verv narrow 
and lining the rim, valves short and enclosed fpl. iv. fig, 7). 
Flinders Range near Ilorrocks Pass and Quorn, 

44. Eucalyptus hkmipeiloia E. Muell. ex. Benth. in FI, Austral., 3, 216 (1866), 

sensu stricio. 

Tree. Mature leaves alternate, petioiate, lanceolate to broadly lanceolate, 
7-14 x 1-3-5 cm. Umbels in panicles or in panicle-like groups on leafless portions 
Of the branchlets, 4-8-flowcred. Buds cylindrical-clavate, pedicellate. 7-10 X 
5-6 mm., pedicels subangular and sometimes' the striations pass up on to the tube. 
Operculum conical, smooth, acute or pointed, about as lung as tube or shorter. 
Fruit pynform urceolate or cylindrical, pedicellate, constricted at the orifice, 
6-S x 4-6 mm.; disc small and oblique or lining the top of the tube, the old 
stamina! ring forming a narrow rim to the orifice: valves deeply enclosed Ctrl iv. 
%. 10). ' w 


Maiden notes (Crit. Rev., II, 14) that Mueller in hfe Census gave Fragtn. 
ii, 62 (1860-61) as the reference for the description of this species. However, the 
remarks in this volume of the Fragments hardly constitute a real description, and 
Blakely was tight in giving the reference as above. On the other hand, Blakely 
gives E. albens Miq. in Ned. Kruidk. ArchicL, 4, 138 (1856), despite the fact 
that Mueller in the Eucalyptographia (also quoted by Maiden) points out that the 
name, which was a misprint for E. pallens DC, was given without diagnosis, hence 
the correct .reference should he fi. albens Mi<]. ex Benth.. Fl, Aust., 3, 219 (18661. 
Blakely accepted E. hemiphloia, E. albens and R, microcarpa- as distinct species, 
but the writer is in agreement with Maiden when he considered the last two to 
be varieties of the first, a decision he later revised. As noted by Blakely 
(Key to Euc, 236), "En Victoria it (i.e., E. microcarpa) seems to pass gradually 
into the typical form (i.e., E. hemiphhia),** This is typical of the problem facing 
the student of the genus. Specimens which appear absolutely distinct are found 
on more extensive collecting to grade into one another. £. albens differs in its 
usually larger buds and fruits and in the glaucousncss of the leaves, branchlcts, 
buds and fruits, in no species of Eucalyptus does glaueousness prove to be a 
reliable character, and a study of the dimensions given for E, hemiphlaia and 
E> albens in the Key and hy Maiden (Crit. Rcv. r 58, 443) will show that there is 
a certain overlapping. 

Southern Flinders Range between [.aura and Mount Remarkable. 

44a. Eucalyptus hfmtphloia var. microcarpa Maiden in Crit. Rev., 47, 207 
(1921) ; £. microcarpa Maiden in Crit. Rev., 58* 483 (1923). 

Differs from the above mainly in its smaller buds and fruits which are 
pedicellate to sessile. Buds 6-7 x 3-4 mm. Operculum has an angular tip (also 
seen in species of the odor at a complex). Tube subangular, Eruit cylindrical nr 
.slightly smaller at each end. 4-7 x 4-5 mm. (pi, iv, fig. 11). 

Distribution as above. 

44b. Eucalyptus hemiphlcua var. albens F. Muell. ex Maiden iu Crit. "Rev., 
11, 20 (1914) ; E. albens Miq. ex Benth. in FL Aust,, 3, 219 (1866). 

Differs from the typical form in being glaucous on all parts and in the usually 
larger buds and fruits. Blakely gives the following dimensions : Suds 10-15 x 
5-8 mm. fruit 9-12 x 8-10 mm. In most of the South Australian material these 
organs arc smaller but still larger than the dimensions for E.hemtptdoia stnsu 
stricto (pi. iv, fig. 9). 

Distribution as above. 

45. Eucalyptus largtflorlxs F. Muell. in Trans. Vict Inst., 1, 54 (1855); 

E. bicolor (A. Cunn.) Hooker in Mitchell's lour. Trop. Aust., 390 

Tree. Mature leaves alternate, petiolate, lanceolate, surface dull and faintly 
bluish, 7-^3 x 1-2 em. U-ntbch' in axillary or terminal panicles* 3-8-flowered, 
peduncles slender. Buds obovoid or davate on slender pedicels, 3-5 x 2-3 mm. 
Operculum hemispherical or almost conical, shorter than tube. Fruit cylindrical 
or pyriform, the orihee slightly constricted, pedicellate, 3-4 x 3-4 mm.; rh'.sc very 
narrow, valves enclosed and short. 

Along the River Murray. Also occasionally in the Flinders Range. 

46. Eucalyptus Behrtaka F. Muell* in Trans. Vict. Tnst, 1, 34 (1855). 

Malice or small tree. Mature leaves alternate, petiolate. ovate or broadly lanceo- 
late, 7-12 x 1 "5-5 '5 cm, Umhtds 3-6-llowcred, in terminal panicles which may be 
open or short and compact. Buds sessile, cylindrical to obovoid, 5-6 x 2-3 mm. 
Operculum hemispherical or shortly and obtusely conical. Eruit sessile or almost 


so, hemispherical or obovoid, 3-5 x 3-5 mm.: disc narrow, valves short, enclosed 
or just below the orifice. 

Blakely gives 5-7 mm. as the length of the fruit, hut none as large are present 
IH South Australian specimens. Maiden figures (Crit. Rev., pi, 48, fig. 6) some 
larger fruits, but the disc is quite different from that in other material oi the 
species and the identity U therefore in doubt, 

Eyre Peninsula, southern Flinders Range, Murray Desert and Kangaroo 

47. Eucalyptus fasciculosa J\ Muell. in Trans. Vict, Inst., 1, 34 (1855). 

Medium-sized tree. Mature leaves alternate, petiolale, lanceolate to broadly 
lanceolate, 8-18 x r 5-3*5 cm. Umbels in axillary or terminal panicles. Buds 
clavate on sttbangular pedicels, 4-6 x 3-4 mm. (pedicels 3-5 mm. long'), usually 
with numerous oil glands. Operculum shortly conical and narrower than top of 
tube. Fruit pedicellate, obconical, cylindrical or pyriform with a thin, rim lined 
by the disc, valves short and enclosed (pi. i, fig. 6). 

Mount Lofty Range to Kncounter Ray, Murray Bridge, Upper South-East. 

48. Eucalyptus cladocalyx F. Muell, in Litmaea, 15, 388 (1852). 

Small to medmm-sized tree. Mature leaves alternate, petiolate, lanceolate to 
broadly lanceolate, 8-14 x 1*3-3 cm. Umbels clustered on leafless portions of the 
stems or in panicles, peduncles terete. Buds cylindrical, urceolate, pedicellate. 
10 x 4-6 mm. Operculum hemispherical, apiculate, much shorter than tube but 
sometimes a little wider, 3 x 4-6 mm. Fruit ovoid or cylindrical-urceolate, having 
a constricted orifice, almost smooth or becoming rugose or costate on drying, 
10-15 x 5-10 mm. ; disc forming a narrow rim, valves deeply enclosed (pi, i, fig. 5}. 

Eyre Peninsula, southern Flinders Range and Kangaroo Island. 

49. Eucalyptus gamopiiylla F. Muell. in Fragtn.. xi. 40 (1878). 

Glaucous malice. Mature leaves opposite, sessile or ampiexicauk, orbicular, 
lanceolate or cuneate when the pairs are joined, 5-8 x 1-2 cm. Utnbel.s poorly 
formed, singly or in short panicles, in the upper axils or terminal. Individual 
umbels with few flowers. Buds clavate or pyriform, pedicellate, 5-7 x 4-6 mm. 
Operculum hemispherical, much shorter than the obconical or pyriform tube. 
Fruit campanulate, pyriform to almost urceolate, 8-11 x 6-8 mm.; rim sharp 
around the narrow oblique disc, valves short, broad and triangular, near orifice 
or enclosed. 

Blakely slates that this is :t very glaucous species, but our material was only 
iuft glaucous. This character is never very dependable. 

Near the north-western corner of the State. 

Species Not Included in the Kev 

There is r group of tropical species which is known to extend into South 
Australia. Material of these is by no means satisfactory, since buds, fruits and 
leaves arc rarely included on the same specimen and notes regarding habit and bark 
character are usually lacking. Consequently there is a certain doubt concerning 
the true identity of the specimens, and the species have, therefore, been omitted 
from the key. However, an account of the South Australian Eucalypts would be 
incomplete without some reference to them. 

In his Flora of South Australia, Black includes E* pyrophora Benth. and 
E. ierminalis F. Muell. (p. 420), and in his additional notes (p. 694) it is stated 
that the White Wash Gum of Central Australia is £. papuana F. Muell., not 
£. terminalis F Muell. In a later paper, by J, B, Cleland, for which Black identi- 
fied the plants (6), there is a reference to E, dichromophlola F. Muell., which has 


been recorded for the Mann and Musgrave Ranges by Black (3), and, in the 
same journal, a paper by Blakely (4) describes E. papimna var. aparrerinja, In 
the Key, Blakely lists R. femiginea F. Muell. as South Australian but gives a 
Ceutral Australian locality. 

All these species come in the Corymbosae and all belong to the Macantherae. 
Without adequate material and field notes it is difficult to distinguish between 
some of them. Until further collecting is done it will be. difficult to decide which 
of them should be recorded for this State. 

Another species of interest is E, gongylocarpa Blakely (4), for which one 
South Australian specimen is recorded with the type description. This came from 
"25 miles S.S.W. of Mount Watson/' but consists of leaves only and, in Blakcly's 
opinion, represents the juvenile slate of the species. This requires checking. 
2f. trivdva Blakely, I.e., and E, orbifolia F. Muell. (4) (7) may also occur in 
this State. 

& Kalangadooensis Maiden and Blakely is described as a probable hybrid. 
The type material appears to represent an unusually large form of E. vhninalis 
var. Huheriam. Further collecting may determine the status of this form. 


The writer wishes to acknowledge the financial assistance given by the 
authorities of the University of Adelaide, which made possible a visit to the 
important collections in the National Herbaria at Melbourne and Sydney. 
Reference must also be made to the helpful co-operation of the staffs of these 
institutions, both during the visits and at other times. 

Mr. J . M. Black and Professor J. B. Cleiand kindly lent their valuable collec- 
tions, which include a number of fype and other important specimens. The 
Eucalypts among the specimens of the Tate and the Elder Exploring Expedition 
Collections were also consulted in the Flerbariurn of the Botany Department of the 
University of Adelaide. 


(1) Bentham, G. 1866 Flora Australiensis, 3 

(2) Black, J. M. 1929 Flora of South Australia, Govt. Printer, Adelaide 

(3) Black, J. M. 1934 Trans. Roy. Soc. S. Aust., 58, 168 

(4) Blakely, W. F. 1936 Trans. Roy. Soc. S. Aust., 60, 153 

(5) Brett, R. G. 1937 Papers and Proc. Roy. Soc. Tas. ( 75 

(6) Cleland, J. B. 1936 Trans, Roy. Soc. S. Aust., 60, 114 

(7) Gakdkek, C. A. 1940-41 Jour, Roy. Soc. W. Aust., 27, 187 

(8) Jessup, R. W. 1946 Trans. Roy. Soc. S. Aosl, 70, 3 

(9) Mueller, F. von 1879 Eucalyptographia 

(10) Worm, J. G, 1937 The Vegetation of South Australia, Govt. Printer, 

Frequent reference is also made to Blakely's "Key to the Eucalypts" and to 
Maiden's "Critical Revision of the Genus Eucalyptus/' 


Plate I 

1 £, axytmtra (from type in Tate Herb.) 5. E. dadocalyx 

2. E, pyriformis & E. fasctculosa 

3, E. pachyphylld £■ £ calciadfrxx 

4 E Mvrrisii 8 - £ - Ewartuma (from portion of type in 

J, M, Black's collection.) 

(All drawings natural size.) 


1. E, remota 

2. E. oblique var. megacarpa 

3. E. d-iversifoUa 

4. E, leucoxylcm var, paupertta 

5. E. leutaxylan- var. pauperita 

6. E. leucoxyton var. macrocarpa 

Plate II 

7. E. Icucoxylon sensu stricfo 

8. E. incrassata sensu striclo 

9. E. incrassata snisu stricto 

10. E. incrassata var. costala 

11. E. incrassata var. angulasa 

(All drawings natural size.) 


1. £. Baxter* 

2> E. viminalis var. Ilubcriana 

3. E. ovata 

4. K. tlaeophora 

5. £. eosinophylic* 

Plate TTI 

6. & plnipinia-na 

7. E. conglohata 

8. _£"„ wiceps 

9. £. dutnosa 
10. £. rugasa 

(All drawings natural size.) 

11. £. pileata 

12. i£. &rac%fa^v^r 

13. Hi ca»anwa 

14. JEs, gra-cilis 

15. £. culycoyona 


1. E. oleosa var. f/Iauca 

2. E. oleosa var, angustifolia 

3. E. Flocktoniae 

4. E. oleosa sensu stricto 

5. E* odorata 

6. E. odorata var. aligns Itfolia 

7. E. viridis 

Plate IV 

8. E. La-nsdowneana 

9. E. hemiphloia var. albens 

10. E. hemtphloia sensu stricto 

11. jB. hemiphloia var. microcarpa 

12. £. leptopkyllat 

13. £. ttMciwafo 

14. It. inter iexta 

(All drawings natural size.) 


By C, G. Stephens 


Soil is composed of weathered rock material modified by additions of organic matter derived from 
vegetable and animal life. Despite the obvious association and contributions of rock, climate and 
organic matter, there have been in relatively modern times different genetical classifications of soils 
which have rested purely on geological, climatic or organic factors of soil formation. Largely 
because of the language difficulty these monogenetic classifications were but slightly influenced by 
the work and theories of Dokuchaiev until well after the beginning of this century, four or five 
decades since his earlier publications on the genesis of soils. There seems little doubt that 
Dokuchaiev was the first to fully realize the function of more than one factor in soil formation, and 
that the factors were not independent of each other. In this connection Neustruev (1927) states: 
"., should be noted that though Dokuchaiev had established such factors of soil formation as 
climate, parent rock, relief, vegetation and age of the country, he still, in the first place, emphasised 
the idea that it is their interaction which leads to definite conditions of soil formation." Ototzky 
(1946) quote Dokuchaiev as follows: "Knowing all the physico-geographical elements of a certain 
locality, it is easy (!) to predict which soil covers it ." 



By C. G. Stephens* 

[Read 10 April 1947] 

Soil is composed of weathered rock material modified by additions of organic 
matter derived from vegetable and animal lite. Despite the obvious association 
and contributions of rock, climate and organic matter, there have been in relatively 
modern times different genet! cal classifications of soils which have rested purely 
on geological, climatic or organic factors of soil formation. Largely because of 
the language difficulty these monogenetic classifications were but slightly 
influenced by the work and theories of Dokuchaiev until well after the beginning 
of this century, four or five decades since his earlier publications on the genesis 
of soils. There seems little doubt that Dokuchaiev was the first to fully realize 
the function of more than one factor in soil formation, and that the factor's were 
not independent of each other. In this connection Neustruev (1927) states; 
". ... it should be noted that though Dokuchaiev had established such factors of 
soil formation as climate, parent rock, relief, vegetation and age of the country, 
he still, in the first place, emphasised the idea that it is their interaction which 
leads to definite conditions of soil formation." Ototzk3 f (1946) quotes Doku- 
chaiev as follows: ''Knowing all the physico-geographical elements of a certain 
locality, it is easy (!) to predict which soil covers it." 

Dokuchaiev gave the formula 

P=f (KO,C,V), 
in which P stands for soil (pochva), K for climate (klimat), O for organisms 
(organism), G for subsoils (gornaya poroda) and V for age (vosrast) of the soil. 

According to Neustruev it was Sibirtsev who first established the frequently 
"predominant role of climate," "the idea of zonality of soils and zonal types of 
soil formation.' Sibirtsev also distinguished "a division of intrazonal soils" 
to which,, in addition, he applied the term semi-zonal, thus acknowledging their 
partial dependence "on climate and their occurrence m definite zones beyond 
which they are found hut rarely, and that in peculiar conditions." 

A recent mathematical treatment of the role of factors in soil genesis is that 
given by Jenny (1941) in his book, "Factors of Soil Formation." He expresses 
the relationship between soil properties (and hence the soil which is an ensemble 
of soil properties) and the environment by two equations: 

$~& (cl'.o'.r'^t), 
and s = f {cl, o, r. p, t ) 

in which the symbols el'. o' t r\ p and t represent soil climate, soil organisms, 
soil relief, parent material and time, and el, o and r represent environmental 
climate, organisms* and relief; s is any soil property, and the soil-forming factors 
represented by the symbols are postulated as independent variables, i.e., "ihey are 
the independent variables that define the soil system." 

From the second of the above equations Jenny derives an equation of partial 
differentials, namely; 

3s\ / ds 


te= (—) <k\+ I—) do 4- L] 

V del / fl. r P £>, t. * do' cl, r, p, t. \ dr/ cl, o, p, t. 

+ LJ\ dp + £_] d 

\ dp/ cl, o. r, t. \ 9t / cl, o, r. p. 

*Wa»te Agricultural Research Institute, University of Adelaide, 
Trans. Roy, Soc, S. Anst., 71, (2), 1 December, 1947 


The interpretation of this equation is that the total change of any soil pro- 
perty, and hence differences between soils, depends on and is a function of the 
sum of the changes of the soil-forming factors * also, assuming that it is possible 
to assign a numerical scale to each of the variables, the magnitudes of the partial 
differentials arc true indices of the. importance of the various factors. 

In reality it is not possible to derive the values of the partial differentials 
because of actual and uncontrollable variation in one or more of the remaining 
variables, and because it is extremely difficult, if not impossible, to assign a 
numerical scale to some of the variables, particularly o and p. In addition, cl, o, 
r, and p are, or can be, multiple factors and yield groups of functions; for 
example, the ratio of precipitation to evaporation (P/E) and temperature (T) 
arc largely used to characterise ci. Furthermore, the variables, despite 
their being assigned an independent status to facilitate the mathematical 
approach, arc in nature not completely independent, relief for instance, as is 
well known to gcomorphologists, being a function of parent material and climate, 
but having a partially independent relationship with soil. The treatment is thus 
to a degree philosophical, rather than rigidly mathematical. Hence it is necessary 
to work out the significance and role of the various soil-forming factors on a 
statistical, graphical, or diagrammatic basis. 

Jenny illustrates the function of each of the soil-forming factors by dis- 
cussing them in turn in their relationships to various soil characteristics. He does 
not attempt to show the effects of interaction or illustrate the variables operating 
together; rather his efforts arc directed towards separating and illustrating the 
effects of the individual variables. 

More recent work by Jenny (1946) illustrates the relationships of soil series 
and types mapped in various soil surveys to the individual soil-forming factors, 
particularly how various sequences of soils are related to changes in the value 
or composition of one of the variables. lie postulates ''five canonical functions 
of pedology 17 where four of the variables remain constant, thus: 

Climof unctions - - s= f (cl) r p t 

Biof unctions - - s = f (o) c j r p t 

Topofunctions - " s == ^ ( r ) cl o p t 

Li thof unctions - - s = f(p)clort 

Chronofunctions - - s = f (t) c j Q r 

Designating an ensemble of s values by E(s) he writes: 

Soil es E(s) = f (cl, o, r, p, I ) 

and defines the various soil sequences as follows: 

Clunosequences - E(s) = f (d) o r p t 

Bioscquences - E(s) = f (o) C J r p * 

Toposcquenccs - E(s) =f (r) c | p ^ 

Lithosequences - E(s) = f (p) c i r ^ 

Chronosequences E(s) = £ (t) c ^ r 

Toposequcnccs include clinosequences and hydrosequences, the former cover- 
ing the effect of the slope factor (i) and the latter the effect of the water table 
factor (w) on soil formation. 

Each of the soil sequences is illustrated by reference to maps and diagrams 
relating to soil series defined in soil surveys. Examples quoted are as follows: 


Chronosequencc: (Tujunga), Hanford, Greenfield, Ramono, Placentia, 

(San Joaquin) series. 
Lithosequence: Colma, Hugo, Sweeney, Sheridan series. 
Toposeqitences: See Clinosequcnce and Hydrosequence, 
Clinosequence: Sheridan sandy loam, shallow phase, Sheridan sandy 

loam, deep phase* 
Hydrosequence; Panochc clay, Oxalis clay, Levis clay. 
Clmwsequence : Panoche, Sorrento series. 
Biosequence : Parr, Octagon, Miami series, 
Jenny states that every homogeneous soil type or series belongs theoretically 
to each of the five canonical functions and Illustrates this by pointing out that 
the Panoche series is a recognisable member of more than one soil sequence. He 
thus indicates a method of recognising the role of more than one variable in 
determining the soil series or type. 

Recently, in Wilde's "Forest Soils and Forest Growth" (1946), Dokuchaiev 
was quoted as follows : "The soil is a result of reactions and reciprocal influences 
of parent rock, climate, topography, plants, animals and age of the land." Wilde's 
formulation following this is 

s=r / &£*9 dt 

in which S is the soil, g is geological substratum, e is environmental influences, 
b is biological activity, and t is time. This equating of the soil as an integral of 
the soil-forming factors against time has a distinct appeal as a more precise 
formulation that that of Dokuchaiev or that of Jenny. In particular the value 
of some of the variables has changed with the passage of time: for instance 
certain soils largely owe their morphological characteristics to a climate operating 
in past geological time, e.g., lateritic soils were formed in Australia most probably 
in the Pliocene, and as they exist in "'fossil*' form today are not particularly 
affected by or responsive to the prevailing climate. Wilde apparently docs not 
regard g, e and b as independent variables. 

The purpose of the paper presented here is to record a diagrammatic method 
of synthesis of the role of the soil-forming factors and to relate the components 
of the diagrams both to the independent variables of the Jenny equations and to 
the partially dependent and partially independent variables which the soil- 
forming factors actually comprise in nature. 

Let the variables which determine the soil (S) be represented as follows: 

C for climate 

O for organisms 

R for shape of the land surface 

W for height and other features of the water-table 

P for parent material 

T for time 

In nature these are not completely independent variables and their known 
dependencies are set out in Table L 

Table I 
A change in C may cause a change in O, R t W* P, S but not in T 

tt t> » v t> n ft 7} >i *— » VVj t I K> , f ,, „ 1 , K. 

t\ h tt *S >t j» >» }> >» ^- j j v4i Wj * , 3 ,, n tJ X 

t' « jj ** i» n 15 »j if U, JT, O „ f , tr L t \^ t i\. 

«j it Jt ■*■ n m i» ri >« tJ, K., W, O ,, ,, t, 1} (J 

it tt ii «*■ it ti »• *» s« V_t L/j iv, W, JT f J 

.- h „ S „ „ , t „ „ O, R, W, P but not in T, C 


A complete list of the possible relationships between any two of the variables 
Is given by the partial differentials in Table II. 

Table II 


as ds as as as as 

3C 30 3R 3W 9P 3T 

ac ac ac ac dc ac 

as do dR aw ap bt 

ao do ao do ao ao 

as ac br aw ap st 

3R 8R 8R 3R 8R 8R 

as ac ao aw ap 3t 

aw aw aw aw aw aw 

as ac ao aR ap aT 

ap ap ap ap ap ap 

as ac ao aR aw 8t 

3T 6T 3T 8T 3T 3T 

as ac ao 3r aw ap 

Reference to Table 1 and a consideration of the lack of effect of changes in 

3T 3T 3T 3T 3T 3T 
C O, R, W, P and S on the change of Tshovv that > > * > & 

as ac ao 3r aw ap 

are of no consequence in soil formation and that T is a completely independent 

ec ac aR 3R 

variable. Similarly , , , & do not operate. Hence the relationship 

aw ap ao aw 

between a soil.and the soil forming factors is probably best expressed like Wilde's 

equation as: S = J f (C, O, R, W, P) dT 

in which T is independent and C, O, R, W and P can have both dependent and 
independent status. 

The function of these soil-forming factors can be qualitatively (and some- 
times quantitatively) but precisely indicated both as independent and interacting 
variables by the diagrammatic and mathematical representation of the relation- 
ships of the soils mapped in soil surveys to the variations in the factors which 
characterise the environment of the area surveyed. Provided the environment 
is studied in the necessary detail during the course of a soil survey, it is possible 
to determine in some degree the relative importance of the different soil-forming 
factors in the genesis of the different soil series or types. This has been done in 
some soil surveys in Australia, with the result that it is now possible to present 
useful diagrammatic expressions of the dependence and interaction of the soil- 


forming factors. The comparison of a soil map based strictly and entirely on 
field observation of soil morphology and distribution with topographic, hydro- 
graphic, climatic, geological and vegetation map* and a study of the Tertiary and 
Recent geomorphology of the same area are generally most revealing regarding 
the functions of climate, organisms, relief, water-fahle, parent material and time 
in soil formation. 

The construction of diagrams to illustrate the importance of the factors and 
the order in which they have operated calls for some ingenuity, but it has been 
found possible in a relatively simple arrangement to integrate the functions o£ 
the variables operating in the formation of different soil scries and types occurring 
in the same locality. Examples of three such diagrams relating to widely 
separated surveys of various sizes are illustrated in fig. 1, 2 and 3. Two of these 
diagrams, in a provisional and rudimentary form, and devoid of any mathematical 
interpretation, were published previously in the bulletins describing the relevant 
soil surveys; namely in: 

1. CS.LR. (Australia) Bulletin No. 150, 1942, 'The Soils of the Parishes 
of Longford, Cressy, and Lawrence, County Westmorland, Tasmania." 

2. CS.I.R. (Australia) Bulletin No. 188, 1945. "A Soil, land-U^e and 
Erosion Survey of Part of County Victoria, South Australia." 

The third diagram relates to the following: 

3. Transactions of the Royal Society of South Australia, 67, (2), 1943, 191- 

199, "The Pedology of a South Australian Fen." 

In iheir present form the diagrams give qualitative expression to the variables 
of the Jenny equations and provide a means of recognition of his soil sequences, 
which arc naturally segregated in different portions of the diagrams. They also 
confirm the crude additive character of the soil-forming factors expressed bv his 
equation of partial differentials. In addition they provide a framework or 
reference grid on which can be recognised the appropriate place at which the 
partial differentials of the partially dependent and interacting variables 
C O, R, W, and P operate in determining the course of soil development, in 
relation to the independent variable T. Inspection of the diagrams shows how 
a "lateral analysis" relates the various steps in soil formation to the independent 
Jenny variahles and how a 'Vertical (time) analysis'' reveals the dependent 
reaction of the natural variables and their ultimate control of soil formation. The 
diagrams should provide a useful starting point in the quantitative evaluation of 
the factors C R. W and T, which lend themselves to such treatment. 

Fig. 1 is a pedogenetic diagram of the soils of the eastern portion of County 
Victoria in the wheat-growing and sheep-raising country of South Australia, It 
includes all of the Boils mapped over an area of about 600 square miles, showing 
how each soil series is a synthesis of the functions of the variables dealt with 
above. The morphology of the soils, a soil map, a topographical map and 
diagram, a rainfall map and other climatic data and details of the geology and 
vegetation of the area were published in the soil survey bulletin mentioned pre- 
viously. In addition, in this diagram the contribution of calcareous loess to the 
morphology of two of the soil series is shown: this following the recent work 
of Crocker (1946). 

in the first column on the right-hand side of the diagram is shown the 
relationship of the various steps of the diagram to the Jenny variables. Recogni- 
tion of his soil sequences is easy, for example the Calto*ne, Canowie and Bunda- 
leer series comprising a cltraosequence. In addition, soils whose genesis is 
dominated by one or more variable? can be recognised. For example, both the 
Yangya silty-loam and the Beetaloo series are formed on calcareous materials in 






i ■ 



a u 


4 © « 















3 a 

1 1 


g a 

; a 

! I 

I I 

| o 

i § 


all localities in the area, irrespective of the change ol the leaching factor under 
a change of rainfall from 14 to 24 inches per annum; i.e., the morphological 
characters of the two soils are predominantly governed by the geological factors 
relief and parent material, climatic variation in the region surveyed producing 
no correspondingly significant variation in profile characteristics. * However, on 
passing further into the adjacent lower rainfall country, the climatic factor 
operating on this parent material does become significant, for at rainfall levels 
of 12 inches per annum and less a soil devoid of the nearly black sub-surface 
layer characteristic of the Yangya silty loam is formed. Such a soil has been 
described at Melton by Prescott and Skewes (1938). 

Soils dominated by the climatic factor are the Caltowic, Canowie and Bunda- 
leer series, and, corresponding to approximately the same climatic variation but 
on different parent material, the Pirie, Yarcowie and Belalie series. The Caltowic, 
Yarcowie, Canowie, Belalie and Rundalcer series comprise the malice, red-brown 
earth and podsolic zonal soils of the region. Retention of calcareous loess in the 
Caltowie and Yarcowie series, as well as being a function of climate, is also 
governed by relief, and similarly both variables operate in the accumulation oi 
cyclic salt in the Piric series. 

There are two aspects of time as a soil-forming factor which are important.. 
First, as used in the body of the diagrams, lapse of time since initiation of soil 
formation is a real measure of actual age of the soil. However, such a time 
lapse does not necessarily precisely specify the maturity aspect of soil develop- 
ment, since maturity can be reached at various rates. Maturity is a characteristic 
of the soil itself and should be regarded as a dependent variable comprising one 
of the S values which make up the soil ensemble. In the diagrams the soils are 
classified as immature, mature, over-mature and senescent. Concerning the 
classification of the solonchak— the Piric series— the designation as immature is 
in conformity with the possible development of this soil to a solonetz ami 
ultimately a solod, but since it is in equilibrium with the present accession of 
cyclic salt, it may equally well be considered as mature. 

^ In the second column on the right hand side and at appropriate places in the 
diagram partial differentials indicate differentiation of certain of the variable* 
witrt respect to change of other variables. Fur example, the establishment of 
different parent materials- in the various sites where soil formation ensues is 

3R 5R ap dV 
shown to he dependent on the partial differentials . „ . ivhile the 

f b dP dC dl< dC 

effect at climate and organisms (restricted to vegetation in the diagram) is 

dO dS dS 

indicated by the differentials , & , The Junctions of this latter group 

ac so ac K 

of differentials is dealt with more comprehensively following the discussion of 

Fig, 2 h a pedogenetic diagram of the soils of the Longf ord-Cressy district 
which lies in the Launceston [Tertiary Basin in northern Tasmania. It relates to 
all the soils mapped over an area of approximately 100 square miles in a region 
devoted to mixed farming. The details of the morphology of the soils, a soil 
map, a cross sectional diagram showing the relationshio of the soil to topo- 
graphical and geological features and climatic and vegetation data were published 
in the soil survey bulletin referred to above. 


2*3 3 

-4 -J 9 



o * 




As in fig. 1 the relationship of the various steps of soil formation to the 
independent variables of the Jenny equation is shown. Also the function of the 
partial differentials is illustrated by the upper portion of the diagram involving 

3R 6R dv ap ac to as as 

, 1 & ant j the lower , , & 

dv dc 6r ar ax ac ao ac 

This diagram is of particular interest because of the age of some of the soils, 
i.e., time varies widely, and consequently there has been a correspondingly large 
variation in the climate to which some of the soils have been subject. Hence, 
unlike fig. 1, where geographical variation in climate was of the greatest signi- 
ficance, this diagram refers to a restricted area of considerable chronological 
variation of that function and its corresponding effect on the development of the 
older soils- Consequently some of the soils are senescent. 

It is appropriate to point out at this point that since podzolisation is largely 
an irreversible process, and since there is a variation of climate with passage of 
time, the soil will not fully respond to the lowered leaching factor (regrade) in 
drier periods. In fact, further soil development is probably only induced when 
the leaching factor exceeds a value near its previous maximum. Hence the 

influence of on soil formation is largely as a discontinuous integral function. 

_ In fig. 1 and 2 the function o[ the organisms variable has been only partially 
indicated, dependence of the soil, both on the vegetation and directly on the 
climate being indicated. Fig. 4 shows for one soil, the Woodstock sand, a more 
comprehensive treatment of the operation of the time, climate, organism and soil 
variables and indicates the dependent reaction of climate on time and of 
organisms tmd soil on climate with an interaction between organism and soil. If 
consideration of O is restricted to the vegetation, this type of diagram if applied 
to each soil provides an ecogenetic treatment as well as' a pedogenetic one. Jt is 

ac as as as as 

apparent that soil is a function of , , & . covers the 

3t ac ao dr st 

alteration of the soil for constant values of C and O, that is, maturation 
under a steady environment. Vegetation on the other hand is a function of 

ac ao ad ao 

, , & the last term including the effects of evolution, i,e. in this 

ax ac as ax 

case the development and extinction of species in the area since Pliocene times 
where such genesis has been independent of the environment. 

The Eucalyptus satkifolia woodland associated with (lie Woodstock sand is 

l ,. - ,. 3° SO 
an edapho-chrrrahc climax: hence $ have plaved co-dominant roles 

ac 3S ao 

ifl its formation. In the case of a climatic complex pinvs the more prominent 


pan and the same vegetation association mav extend over a number of soil tvues* 
dO ' yv i 

ior an edaphic climax is the more significant function 



m n 
s o 

j3 i3 ** 

" 3 l 

O (h a 

*> 4 « 

P. >. 

3 t* 2 









] f 


5 53 









o to 
id *m 

s : i 


--. *i 

w> ■•> « 


• *. 

C 4 « 


O -i 

•n O C 


S. >H 

^ O -1 

B ,-1 r* 








« r-l 


» ** 

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.o — 

* M 



b im a 

— •id 




o O 


+-. rH 




a O 

« H 


Bj dl 


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5 J 



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> .*' 

■H * 






9 L . 

*3 43 

x: a 


























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o f- 


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;•. ,'.'.■] <h 


o | -■* 


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0) h 




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o x> 






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O <M 






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. e ... 

P! C 




4> -i 





P0 ■« 








+j -. M 

*> Ml-- 


The third figure, is a pcdogcnetic diagram of the soils of Eight Mile Creek; 
Swamp, a coastal fen formation in the lower south-east of South Australia. The 
diagram relates to the four named soil types found on the fen proper, which 
covers an area of about 3,500 acres. The details of the morphology of these 
soils, a soil map, a cross-sectional diagram of the fen showing the relationships 
of the soils to very small changes in relief, and climatic and geological data ate 
included in the publication referred to above. In addition, the ecological relation- 
ships are described in the text and figure of a paper on the vegetation of the fen 
by C. M. Eardlcy (1943), who considers the age of this peat formation to be iess 
than 5,500 years. The simplicity of this diagram distinguishes it from the other 
two to a marked degree. This simplicity is due to the dependent sequence of 
events following the establishment of the topographical factor. This determines 
the position of the water-table, which in turn is the selecting factor operating 
on the climatically and geographically available species and thus deiermines the 
plants growing on any location. The plants, in turn, determine the nature of 
the parent materials which, all being of about the same age, give rise respec- 
tively to the four soil types. The establishment of the relief factor is a simple 
process as compared to the geomorphological processes involved in the establish- 
ment of the parent material and relief factors in the two former diagrams. 

As seen on the diagram, it is possible to indicate the relevance of the Jenny 
variables, but quite obviously the variables involved are not independent but 
rather dependent on a fore-runner variable modified in one case by the effect of 
the local climate in determining the total number and type of species available 
to colonise the area. The dependence of the variables is shown clearly by the 
sequence of partial differentials. 

This diagram indicates that the same four soils comprise simultaneously a 
hydrosequence, a biosequence aud a iithosequence. 

Considering along general lines the above diagrams and temporarily retaining 
both a dependent and independent status for T at well as for C, O. R, W & P, 
we may derive from 

Ss=| (C,0,R,W,P,T) 
the following equations of partial differentials. 

. Where T is independent and C, O* R, W and P are functions of T and of some 
other set of independent variables 

DS as 3C 3S 60 3S dR 3S 3W 3S 3P dS 

=. + + H -I- + 

3T dC dT dO 3T 3R 3T 3\V 3T dV dT 3T 

Similarly when P is independent and C, O, R, W and T ntc functions of P 
and of the other set of independent variables 

DS 3S 3C 3S 30 35 3R 3S 3W 3S 3S 3T 

— = 4, 4- — *. — + + — -f- 

3p ac dv ao 3p an ap aw ap ap 3T sp 


Similarly for W, R, O and C in turn 

ds as ec es 90 as 3r as as ai j as aT 

aw ec aw eo aw 3r aw aw ap aw aT aw 

DS as aC dS 50 dS 8S gW dS 3P SS dT 

an ac an ao an ait aw an ap $& a r r aR 

us as ac as as aR as aw as ap as aT 

ac ao ao aR ao aw ao ar ao ax ao 

ds as as ao as B% as aw as ap as aT 
dc dc aoac an ac aw ac ap ac aT dc 

These comprise a set of simultaneous equations which mathematically define 
the soil. Allowance must be made for T being independent so that on causal 

aT aT aT aT a r r 

grounds , , ,' , and 1 mav he reduced to a constant — say K L . 

ac ao aR aw ap 

ac ac aR aR 

'For similar reasons , and 1 may be represented by IC ; and & and I 

aw ap aw ao 

by Ka. The soil, specified by the very complex integral of the above equations, can 
be most simply represented by the sum: 


— + H- — f — + — + — 

ac ao aR aw ap aT 

as /ac ac ac\ as /ao ao ao ao ao\ 

= — — + — + K* * — + — [ — tn---- + — * — 
dc \ao aR st/ ao \dc aR aw dp arj 

as /3r aR aR\ as /aw aw aw aw aw 

aR\ac ap aT/ aw \ac ao aR ap aT 

as /ap ap ap ap ap\ as / 

± — — + — + — + — -t-i+ — + — K 
ap Vac ao aR aw aT/ st \ 

as as 

This expression includes partial differentials such as , etc. corrcv 

dC dO 
ponding to the effects of the independent variables of the Jenny equations. The 
other terms express the dependent relationships between the variables involved in 
soil formation. 

The above equations and diagrams show the involved manner in which the 
soil-forming factors integrate with one another in the pedogenic process. Although 
the diagrams refer to more or less restricted localities, it is obvious that some 
measure of similar interpretation applies to the broader classification of soils such 
as the soil groups, and points to the fad: that it is futile to attempt too simple a 


genetic classification of the morphologically defined great soil groups. Sufficient 
exploration has now been made to leave no doubt that on a morphological basis 
there is a limited number (probably less than fifty) ot great soil groups. The 
relationship of these groups to their genetic factors is, however, not simple. 

Although climate does play a dominant role in the genesis of certain of the 
soil groups, it has been established that the groups do not lie in exactly similar 
climatic zones and the boundaries of adjacent groups interdigiiate to a consider- 
able degree in the different continents. In this regard the senescent and fossil 
character of some soils is important; as an extreme example the lateritic podsol 
soils of southern Australia extend into arid regions and. besides illustrating the 
above point, are a good indication of the largely irreversible character of the soil- 
forming process, only the slightest detectable reversal of lateritic podsol genesis 
being associated with the drier periods of the climatic cycle. It seems, therefore, 
that, if it is essential to have a genetic classification of soils correlated with the 
morphological one, the arrangement will need to be on a multidimensional basis. 
Such a tabulation is not readily constructed. 





Bfosalyptmt gallclfellft 

Xdapho-Cllaatlc Climax 

Woodatock eand 


La.tari.tlc podsol 



Fig. 4' 


Another consequence of the lack of appreciation of the number of variables 
involved in the genesis of even the great soil groups has been the repeated 
attempts to seek near perfect correlation between large scale climatic and soil 
maps. Similar correlations are sought between climatic and vegetation maps. 
On maps of a continental scale, the mapping of soil and vegetation boundaries 
is of necessity based to some considerable degree on imperfect correlations of 
these features with climate, topography and parent material. Subsequent studies 
•of climate frequently point to correlations of certain lines between climatic 
categories with boundaries of soils and vegetation in fallacious interpretation of 
the degree of importance of the climatic factor. It is obvious that until conti- 
nental soil maps can be prepared almost entirely on a morphological basis the 
effect of the ch'watic factor cannot be fully assessed. For that reason the study 
of smaller areas with precise soil mapping, and in which there is a significant 
geographical and/or chronological variation of climate, is of the greatest 


The author gratefully acknowledges the assistance of ( discussions with 
Dr. W. H. Bryan, Department of Geology, University of Queensland, Professor 
J. G, Wood, Botany Department, University of Adelaide; Mr. E. A. Cornish, 
Section of Mathematical Statistics, Council for Scientific and Industrial .Research; 
Mr. \V. H. Maze, Geography Department, University of Sydney; and Mr. R, L. 
Crocker of the Waite Agricultural Research Institute, Adelaide. 


Crock eh, R. L. 1946 'Tost Miocene Climatic and Geologic History and its 
Significance in Relation to the Genesis of the Major Soil Types of South 
Australia." Counc. Sci. Ind, Res. (AusL), Bulletin No. 193 

EARDLEY y C. M. 1943 "An Ecological Study of the Vegetation of Eight Mile 
Creek Swamp, a Natural South Australian Coastal Fen Formation." 
Trans. Roy. Soc. S. Aust, 67, (2), 200-223 

Jenny, H. 1941 "Factors of Soil Formation/' McGraw Hill Book Co. 

Jenny, H. 1946 "Arrangement of Soil Scries and Types according to Func- 
tions of Soil-forming Factors, Soil Science, 61, (5), 375-391 

Neustkuev, S. S. 1927 "Genesis of Soils," Academy of Sciences of the 
U.S.S.R. Russian Penological investigations, III. Leningrad 

Ototzky, P. 1946 Personalia. Professor Dr. Basile Dokautchacv (1846-1903). 
Official Communications of the International Society of Soil Science, 

Pxescott, J. A- T and Skewes, H. R. 1938 "An Examination of some Soils 
from the more Arid Regions of Australia." Trans, Roy. Soc. S. Aust , 
62, 320-341 

Stephens. C. G-, Baldwin, J. G., and IIoskinc, J. S. 1942 "The Soils of the 
Parishes of Longford, QrcBSjf and Lawrence, County Westmorland, 
Tasmania. Counc. Sci. Ind. Res. (Aust.), Bulletin No. 150 

Stephens, C. G. 1943 "The Pedology of a South Australian Fen." Trans. 

Roy. Soc. S. Aust.. 67, (2), 191-199 
Stephens, C. G., et al. 1945 "A Soil, Land-use, and Erosion Survey of Part 

of County Victoria, South Australia. Counc. Sci. Ind. Res. (Aust.), 

Bulletin No. 188 

Wilde, S. A. 1946 "Forest Soils and Forest Growth." Chronica Botanica Co. 


ByH. H. Finlayson 

At the suggestion of Professor J. B. Cleland, whose researches into the alimentation of aboriginal 
man in this country have now entered upon a gravimetric phase, I herewith record the weight of 
some eighty-eight species and sub-species of mammals of Australian occurrence. 



Ry H II. Finlavsox 

[Read 8 May 19471 

At the suggestion of Professor J. ft. Geland, whose researches into the 
alimentation of aboriginal man in this country have now entered upon a gravi- 
metric, phase, I herewith record the weight of some eighty-eight species and sub- 
species of mammals of Australian occurrence. 

The published data on this head is scanty and of doubtful value. Tt has been 
chiefly derived from the statements of bushmen and other rural observers, whose 
approach to the matter is sometimes tinged by that frivolity which, in the 
ichthyological JieUb has become proverbial. Even when the intention is of the 
best, the weights so derived are usually of the "estimated" variety, and though 
supposed ability* in the difficult feat of estimating weights b ottcn a matter of 
pride, some gross errors have been introduced in this way. 

All weights quoted in the sequel have been personally determined by the 
writer in the field with the balance, upon recently killed animals, and are recorded 
in grammes and kilos.* 1 * 

It is not proposed here to discuss fully the scientific value of body weight as 
a character in the descriptive treatment of mammals, except to remark that its 
value and appropriateness in a general account of an animal ts sr> great as to lift it 
almost to the rank of a descriptive necessity. No other quantitative datum conveys 
so easily an impression of general bulk, and in conjunction with linear dimensions 
its graphic value in illustrating differences of body build in different species, or 
of sexual dimorphism in the same species, is very great. 

On the other hand, its almost universal neglect by taxonomists is not with- 
out sound reason, owing, in the first place, to the great difficulty of adequately 
correlating body weight with growth phase without a much more intimate 
acquaintance of the animal in life than falls to most systematic writers. In many 
marsupials also, concordant criteria of maturity as derived from epiphysial 
condition in the skeleton, cranial sutures, and dental phase, do not 
correspond with the maximum values for either linear dimensions or body 
weight, and the latter may either markedly increase or diminish after skeletal or 
dental maturity is attained. A second difficulty arises from the massive error 
introduced into the observed weight by unascertained variations in the weight of 
the stomach contents, particularly in large herbivores; and a third, from the 
influence of climatfc, seasonal and general ecological factors, which cannot be 
brought under statistical control as geographic or other racial variation. 

In the list which follows, specie* have been treated on as wide a geo- 
graphical basis as the Held data allowed, but the use of trinomials has been 
limited for the most part to case* where the differentiation is marked, 
or at least has been fully denned. Several growth stages have been included, 
wherever possible, and the terms immaiure, subaduit, adult, and aged, which 

V) To the main body of the data I have subsequently added approximate values far 
a few species of exceptional interest, where weights tif freshly killed animals were not 
available. These values were obtained by weighing preserved material of known history 
and correcting the result with a modulus of dehydration, extracted from the loss of weight 
undergone by similar specimens weighed fresh in the field and submitted to the same 
process of preservation for the same length of tune. These citations arr diCfereiitiarod as 

Ti«h. Rny. Sue. S. Aus(. : 71, (2). 1 December 10A7 


are appended to each citation, are cho?en after a consideration of the criteria 
named above and arc subject to Lhe limitations there noted, In most cases these 
terms have been further amplified by the addition of the head and body length 
in millimetres. The maximum weight found in the field for the adult of each 
species has always been included in the citation, but this does not necessarily 
mean that much higher weights may not be reached. 

Finally, it is desired to emphasise that though each individual observation 
is accurate, the body of data which is collectively represented by them is of an 
interim and tentative character only, and is here assembled to meet a general 
or popular want, rather than a systematic one. In its present form it is quite 
inadequate for (as an example) the resolution of problems of specific, still less 
of subspecific, identity* 

Fuller data will be provided in monographic treatment of many of the 
species here mentioned, and some already so dealt with may be traced ill the 
list of references. 


Qrnithorhynchits anatinus Shaw 

(a) Olangolah Creek, Otway Peninsula, Victoria; inun. 9 ; H. and B. 311 mm.; 
wt. 900 g. 

(b) Macquarie River, East Tasmania; subad. 9 ; H, and % 382 mm.; wt 
1-36 kg, 

(c) Murray River, South Australia; ad. 8 ; H. and B. 50+mrn.; wt. 275 kg. 

Echidna aculcata Shaw 

(a) Yavan Creek, Tumut district, New South Wales; $ ; H. and B, 470 mm.; 
wt. 272 kg. 

(b) Olangolah Creek, Otway Peninsula, Victoria; imm. 3 ; II. and B. 406 mm.; 
wt. 1-82 kg. 

(c) Heathmcre district, West Victoria; ad. $ ; II. and B. 530 mm.; wt. 5*90 kg. 

(d) Flinders Island, Bass Strait, Tasmania; 9 ; H. and K. 450 mm.; wt. 
3-64 kg. 

(e) Arthur River, West Tasmania; 9 ; H. and li 500 mm.; wt. 4*32 kg. 

(f) Macquarie River, East Tasmania; 2 ; H. and B. 485 mm.; wt. 3*19 kg. 

(g) James Range, Central Australia; S ; H. aud B. 468 mm.; wt. 2*50 kg. 
(h) Kooringa, Mid-north South Australia; wt. 2-61 kg. 

Sminthopsis eras sic audata crassicaudata Gould 

(a) Moutajup, Grampian Range, area, West Victoria; ad. 2 ; H. and B. 
82 mm. ; wt. 1 1 g. 

(b) Murray MaHee, South Australia; ad. 3 ; H. and B. 84 mm.; wt 15 g, 

(c) Ibid.; ad. 2 ; H. and B. 84 mm.; wt. 14 g. 
Smhitho/>sis crassicaudata centralis Thomas (1) 

(a) West of Appamunua. Lower Diauianlina River, South Australia; ad. 9 ; 
J L and B. 83 mm, ; wt, 10 g. 

Sminthopsis leucopus Gray 

(a) ITeathmere, West VicLoria; ad. S ; H. and B. 98 mm. ; wt. 32 g. 


Antechinomys spcnccri Thomas 

(a) East of Erliwunyawunya, Musgrave Range, North-West South Australia; 
imm. 2 ; H. and B. 85 mm.; wt. 14 g. 

Chaetocercus crislicauda Kreilt 

(a) Chundrmna, Everard Range, North-West South Australia; stibad. 9 ; 

II. and B. 133 mm,; wt. 68 g, 
(h) Ibid. ; ad. & ; H. and B. 160 mm.; wt. 122 g. 
(c) South of Koonapandi, Musgrave Range, North-West South Australia; 

subad. 9 ; H. and B. 130 mm. ; wt. 60 g. 

Chaetocercus cristicauda Jiil!ic?-i Thomas (2) 

(a) Coonehcri. Lower Diarnantina River, South Australia; ad. $ ; H. and B. 
190 mm.; wt. 175 g. 

Phascogalc flavipes Waterhouse 

(a) Waitpinga, Fleurieu Peninsula, South Australia; ad. 3 ; H. and B, 
112 mm.; wt. 49 g. 

(b) Ibid. ; ad. ? ; H. and B. 104 mm. ; wt. 24 g. 

(e) Mackenzie Creek. Grampian Range, West Victoria; imm. $ ; H. and B. 
93 mm.; wt. 23*5 g. 

Dasyurus znvverinus Shaw 

fa) Cradle Vallev, North-west Tasmania; subad. 9 ; II. and B. 298 mm.; 
wt. 690 g. 

(b) North Macquarie River, East Tasmania; ad. S ; H. and B. 350 mm.; wt. 
1-37 kg. 

(c) Tooms Lake, East Tasmania; ad. g (melan.) ; H, and B. 355 mm.; 
wt, L37 kg, 

Dasyurus geoffroyi Gould 

(a) Chundrinna, Everard Range, North- West South Australia; ad. 9 ; II. and 
B, 290 mm.; wt. 550 g. 

Dasyurus maadatus Kerr 

(a) Welcome River, North-west Tasmania; imm, 3 ; H. and B. 425 mm.; 
wL 1-60 kg. 

(b) Ibid,; ad. 9 ; II. and B. 564 mm. ; wt. 2-04 kg. 

(c) Cradle Valley, North-west Tasmania; ad. $ ; 11. and B. 500 mm.; 
wt. 2-96 kg. 

Sarcophilus harrissi Boitard 

(a) Welcome River, North-west Tasmania; imm. 9 ; H. and B. 442 mm * 
2-50 kg. 

(b) Arthur River, West Tasmania; imm. $ ; H. and B. 415 mm.; wt. 3'23 kg. 

(c) Ibid.; ad. 2 ; II. and B. 550 mm.; 4'09 kg. 

(d) Ibid.; ad, tf ; H. and B. 615 mm.; 8'67 kg. 
Myrmecobius fasciatus fasciatus Waterhouse 

(a) Narrogin, South-west district Western Australia; ad. 3 ; II. and B. 
242 mm.; wt. 293 g. (approx,). 
Myrmecobius fasciatus rufus Wood Jones (3) 

(a) Oolarinna, Everard Range, South Australia; ad. <? ; H. and B, 262 mm ! 
wt. 347 g. (approx.). 



Ferameles gunni Gray 

(a) Ross, East Tasmania; subad. 9 ; fit, and R. 299 mm.; wt. 450 g. 

(b) Dunkeld, West Victoria; ad. 9 ; IT. and B. 313 mm.; wt. 550 g. 

Isoodon auratns Ramsay 

Pundi, 70 miles west of Everard Range, North-west South Australia; svibad. 
<* . H. and B, 235 mm. ; wt. 260 g. 

Isoodon obesuhts Shaw 

(a) West Sister Island, Bass Strait, Tasmania; ad. & ; H. and R. 338 mm.; 
wt. 1'36 kg. 

(h) Blue. Hill, Bothwel] district, Tasmania; ad. i ; H. and B. 357 mm,; 
wt. 1-36 kg. 

(c) Ibid.; ad. 9 ; H. and % 355 mm.; wt, 1-13 kg. 
Thcdacomys lagotis Reid 

(a) Kings Creek, George Gill Range, Central Australia; ad. 2 ; H. and B. 
,^?>5 mm. ; wt_ 1 '37 kg. 

(b) Eewellina, Musgrave Range, North-west South Australia; ad. 6 ; II. and 
_B. 428 mm.; wt. 1*70 kg/ 

(c) Ibid.; imm. 6 ; H. and B. 268 mm. ; wt. 450 g, 
I'halacomys lagotis sagiita Thomas (4) 

(a) Goyder's Lagoon, Lower Diamantnia River, South Australia ; aged fi ; 
H. and B. 385 mm.; wt. 1*56 kg. 

(b) Ibid.; ad. ? . ; H. and B. 291 mm. ; wt. 660 g. 

(c) Ibid.; imm. 5 ; H, and B. 212 mm. ; wt. 210 g. 
Thcdacomys minor nvisctius Fin!ayson (5) (6) 

(a) Cooncheri, Lower Diamantiua River, South Australia; ad. 3 ; H. and 

B, 241 mm.; wt. 435 g. 
(!>) Ibid.; ad. 9 ; II. and B. 247 mm.; wt. 312 g. 
(c) Ibid.; imm. $ ; 1L and B. 193 mm.; wt. 212 g. 
Chocroptis castanotis Gray 

(a) Macdonnell Range, Central Australia; imm. 9; H. and B. 165 mm.; 
wt. 153 g. (approx. )« 

Notoyyctcs typhlops Stirling 

(a) Ooldea district. South Australia; ad. ? ; II. and R. 114 mm.; wt. 66 g. 

(b) Liddlc's Hills, Basedow Range area. Central Australia; PI. and B. 120 mm.; 
wt. 71 g. (approx.). 

Notoryctcs typhlops caitrinus Thomas 

Stun Creek, Kimbcrley Division, Western Australia; :mm. £ ; H. and B. 
94 mm.; wt. 47 g, (approx.). 

Acrobatcs pygmaeus Shaw (7) 

(a) Coomooboolaroo, Dawson Valley, Queensland; ad. .? ; H. and B. 75 rnm.; 
wt. 14 g. (approx.). 

(b) Fitzroy River, Rockhamplon district, Queensland; ad. ?; H. and B. 
73 mm.; wt. 12 g. (approx.). 



Dromkia concinna Gould 

(a) Ileathmere, West Victoria; ad. 9 ; SI* and B. 91 mm.; wt. 21 "5 g. 

(b) Meningic, Lake Alexandria, South Australia; ad. £ ; H. and B. 81 mm.; 
wt. 19 g. 

(c) TorrensvaiC, Fleurieu Peninsula, South Australia; ad. 9 ; H. and B. 
77 mm.; wt, 16 g. 

Petaurus breinceps Waterhousc (8) 

(a) Penola, South-East district. South Australia; ad. o ; H. and B. 163 mm.; 
wt. 130 g. 

(b) Ibid.; aged 9 ; wt. 90 g. 
Trichosurus vulpecitla Kerr 

(a) Cradle Valley, North-west Tasmania; ad. 2 (grey phase); H. and B. 
495 mm.; wt. 4' 10 kg. 

(b) Ibid.; imm. 8 (grey phase) ; H.. and B. 400 mm. ; wt. 1 '81 kg. 

(c) Gorae, West Victoria; ad. o ; IF and B. 546 mm.; wt. 3*63 kg. 

(d) Heathmere, West Victoria; subad. 9 ; H. and B. 362 rnm. ; wt. 1 -75 kg. 

(e) Pccaree, South-East district of South Australia; ad. $ ; H. and B. 
440 mm.; wt. 3*07 kg. 

([) A garden, Adelaide, South Australia; subad. c? ; H. and B. 405 mm.; 

wt. 1*75 kg. 
(g) Palana, Flinders Island, Bass Strait, Tasmania; ad. 9 ; H. and B. 502 mm.; 

wt. 4-09 kg. 
(h) Choorlabinna, Musgrave Range. North-west South Australia; ad. '& ; 

H. and B. 390 mm. ; wt. I- 37 kg. 
(i) Eewellina. Musgrave Range, Norih-west South Australia; ad. 9 ; H. and 

B. 372 mm.; wt. 1*48 kg. 
(j) Wollara, Basedow Range area, Central Australia; subad. ft ; H. and B. 

313 mm. wL 910 g. 
(k) Thangool, Cailidc Valley, Queensland; subad. 9 ; H. and B. 410 mm, ; wt. 

1-36 kg. (9). 

Trichosurus caninus Ogtlby 

(a) Upper Rvan's Creek, North-east Victoria; ad. 9 ; H. and B. 508 mm.; 
wt. 2*73 kg. 

(b) Ibid,; subad. i ; H. and B. 476 mm.; wt 2*16 kg. 
Pctauroidcs volans vofans Kerr (10) 

(a) Green ITills Forest, Batlow district. New South Wales; ad. 9 ; H. and B. 
440 mm, ; wt. 1"36 kg. 

(b) Ibid,; subad. 6 \ II. and B, 415 mm.; wt. 1-02 kg. 

Pscitdochirus lanir/hwsus Gould 

(a) llatherlcigh. South-East district, South Australia; ad. 3 ; H. and B. 
305 mm. ;' wt. 800 g. 

(b) Ibid.; ad. 9 ; II. and B. 318 mm. ; wt. 800 g. 

(c) Glencorrie, Fleurieu Peninsula, South Australia: imm. <5 ; H. and B. 
161 mm.; wt. 140 g. 

(d) A garden, Adelaide. South Australia; subad. 6 ; H. and B. 270 mm.; 


(e) Williamstown. North Mount Loftv Range, South Australia; ad. $ ; H. and 
B. 300 mm.; wt 700 g. 

(f) Penrice, Lower North rjislrict, South Australia; ad. $ ; H. and B. 
310 mm.; wt. 700 g. 

Pscudochirus cookl Dcsmarcst 

(a) Blue Hil!s } Bothwell, Tasmania; ad. ? ; II. and B. 357 mm.; wt. 1*36 kg. 

(b) Macquarie River. East Tasmania; imm. 3 ; H. and B. 290 mm.; wt. 570 g. 

(c) Ibid.; subad. ? ; H. and B. 325 mm.; wt, 900 g. 
Phascolaretos a'nereits adustus Thomas (11) 

(a) Kariboe Creek. Callide Valley, Queensland; aged 8 ; H. and B. 665 mm. ; 
wt. 6-36 kg. 

(b) Coomooboolaroo, Dawson Valley, Queensland; aged 2 ; H. and B. 
635 mm.; wt. 5-45 kg. 

Phascolaretos cinercus victor Troughton (12) 

(a) French Island, Westernport Bav, Victoria; ad_ & ; H. and B. 830 mm.; 
wt. 14-09 kg. 

(b) Ibid.; ad. 9 ; It and B. 730 mm. ; wt. 8*82 kg. 

(c) Ibid.; imm. $ ; H. nad B. 673 mm.; wt. 7'27 kg. 
Tarsipes rostratus Gervais and Verreaux 

(a) South-west Western Australia; ad. $ ; IT. and B. 78 mm.; wt. 13 g. 

(b) Ibid; ad, ? ; II. and B. 81 mm.; wt- 17 g. (approx.). 

Hypsipryvaiodon vioscliattts Ramsay 

(a) Gorge Range, Ingham district, North Queensland; ad. 6 ) II. and B. 
325 mm. ; wt. 500 g. (approx,). 

Potorous tridactyhis Kerr 

(a) Upper Macquarie River. East Tasmania; ad. £ ; 11. and B. 385 mm.; 
wt 1*81 kg. 

(b) Blue Kills. Bothwell, Tasmania; ad. 9- . H. and g, 350 mm.; wt. U36 kg. 

(c) Ibid.; imin. £ ; H. and B. 305 mm,; wt. 950 g. 

(d) Gorae. West Victoria; imm. g ; H. and B. 305 mm.; wt. 690 g, 
(Y) Ibid.; imm. $ , H. and B. 336 mm.; wt. 950 g. (13). (a > 

Caloprymnus camprsfris Gould (14) (15") 

(a) Cooncheri, Lower Diamamhia River, South Australia; ad. 9 ; H. and B. 
277 mm. ; wt. 1 *06 kg. 

(b) Ibid.; subad. $ ; H. and B. 282 mm. wt 850 g. 

(c) Ibid*; imm, 2 ; II. and B. 163 mm.; wt. 195 g. 
Bcttongia lesucnri Quoy and Gaimard 

(a) Yaringa. south-west of George Gill Range, Central Australia; subad. ? ; 
II. and B. 285 mm.; wt. 910 g. 

(b) Desolation Glen. Rawhnson Range, Western Australia; subad. ? ; II. and 
B, 245 mm. ; wt. 770 g. 

< 2 > The weight formerly quoted (850 g.) W»S a typographic! error. 


Betlongia cnnicuhts Ogilhy 

(a) Macquarie River/East Tasmania; ad. 6 ; II. and K. 325 mm. ; wt. 1-59 kg. 
Aepyprymnus mfescens Gray (16) 

(a) Thangool. Callide Valley, Queensland; ad. $■; H. and B. 383 mm.; 
wt. 2-50 kg. 

(b) Ibid.; subad. $ ; H. and B. 3S7 mm.; wt. 3*19 kg. 
Logorclteslcs hirsittus Gould 

(a) South of Koonauandi. Musgrave Range area, North-west South Australia; 
ad. 9 ; H. and B. 338 mm.; wt. 1-70 kg. 

(b) Ibid.; suhad. £ ; R and B. 325 mm.; wt. 1-59 kg. 

(c) Fundi, Avesl of Everard Range, North-west South Australia; inim. S ; 
1L and B. 200 mm. ; wt. 275 g. 

Lagorchestes conspicillatus Gould 

(a) Mareeni Plain, north of James Range, Central Australia; ad. $ ; H and B, 
400 mm.; wt. 3*00 kg. 

(b) Ibid.; suhad. $ ; H. and B. 400 mm.; wt. 3*00 kg. 
Peirogde penkillata herbcrti Thomas (17) 

(a) Spring Creek, Taroom district, Queensland; ad. i ; H. and B. 535 mm : 
wt. 5*90 kg. 

(b) Ibid.; subad. $ ; H. and B. 515 mm.; wt. 5'00 kg. 
Pctrogalc xanthopus Gray 

(a) Oraparinna, Flinders Range. South Australia; ad. 9 ; hi. and B 570 mm ' 
wt. 7*50 kg. 

(b) Ibid.; suhad. # ; H. and B. 550 mm.; wt. 6*36 kg. 

(c) Mount Norwest, Willouran Hills, South Australia; aged 9 ; H. and B, 
490 mm.; wt. 6*13 kg. 

Feirogalc lateralis Gould (18) 

(a) Erliwunyawunya, Musgrave Range, North-west South Australia; ad. S : 
H. and B. 498 mm.; 5*00 kg. 

(b) King's Creek, George Gill Range, Central Australia; suhad. $ ; II. and B. 
425 mm.; wt. 3 -63 kg. 

(c) Glen Edith, west of James Range, Central Australia; aged 9 j II. and B. 
465 mm.; wt. 3 '50 kg. 

Macropvs (Thylogalc) eugcnii Desmarest 

(a) Rocky River, Kangaroo Island, Souih Australia; inim. $ ; H, and B. 
475 mm.; wt. 4'09 kg, 

(h) Ibid,, subad. 9 ; H. and B. 495 mm.; wt. 5*00 kg. 

(c) Ibid.; ad. S ; H. and B. 600 mm.; wt. 7-95 kg, 

Macropus {Thylogalc) flindcrsi Wood-Jones 

(a) Flinders Island* Eyre Peninsula, South Australia; ad. $ • II. and B. 
545 mm.; wt. 6-13 kg. 

(b) Ibid.; ad. 9 ; H. and B. 462 mm.; wt. 5"00 kg. 

Macropus (Thylogale) billardicri Desmarest 

(a) West Sister Island, Bass Strait, Tasmania; ad. S ; H. and B. 623 mm.; 
wt. 9-09 kg. 


(b) Flinders Island, Bass Strait, Tasmania; ad. 9 II. and E. 570 mm.; 
wt 7-50 kg. 

(c) Ilarcus "River, North-west Tasmania; young ad. 6 ; H. and B* 520 mm.; 
wt. 6-36 kg. 

(d) Macquarie River, East Tasmania; subad. 9; H. and B_ 482 mm.; 
wt. 3*18 kg. 

(e) Blue Hills, Bothweil, Tasmania (central) ; subad. 6" [ H. r.nd B. 427 mm.; 
wt. 2-72 kg. 

(f) Macquarie. River, East Tasmania; imm. $ ; H. and B. 349 mm.; 
wt. 1-36 kg. 

Macropns (ivallabia) ruficollis ruficollis Dcsmarest (19) 

(a) Yavan Creek, Tumut district, New South Wales; imm. 9 ; H and B. 
627 mm.; wt. 8T8 kg. 

(b) Heathmerc, West Victoria; ad, & ; II. and B. 785 mm.; wt. 24-09 kg. 

(c) Mount Abrupt. Grampian Range. Victoria; ad. 9 ; H, and B. 685 mm.; 
wt. 13-63 kg. 

(d) Black Range. South-East district, South Australia; ad. S ; H. and B. 
703 mm.; wt 16-81 kg. 

(e) Furiier, South-East district. South Australia; ad. 2 ; H. and B. 662 mm.; 
wt. 14-09 kg. 

Macrojmx (ivallabia) tuftcPtljS hemietH Waterhouse 

(a) Macquarie River, East Tasmania; imm. 9; II. and B. 545 mm,; 
wt. 5-90 kg. 

(b) Welcome River, North-west Tasmania; imm. 9 ; II. and B. 620 mm.: 
wt. 9-09 kg. 

(c) Palana, Flinders Island, Bass Strait, Tasmania; imm. ' ; H. and B. 
613 nm; wt. 10*67 kg. 

(d) Ibid,; ad. 9 ; H. and B. 655 mm, ; wt. 14-09 kg. 

(e) Stanley Point, Flinders Island, Bass Strait, Tasmania ; ad. 5 ; H. and B. 
770 mm.; wt. 24' 54 kg. 

(f) Arthur River, West Tasmania; ad. ? ; H. and B. 740 mm.; wt. 2272 kg. 
Macropus (ivallabia) ualabaius ualabafus Lesson and Garnot 

(a) Blunt' s Cully; Ryan's Creek, North-east Victoria; aged 6 ; H. and B. 
800 mm.; wf. IS- 18 kg. 

(b) Hillas Brook. Tumut district, New Soutb Wales; ad. 2 ; H. and B. 
760 mm.; wt. 14*79 kg. 

(c) Ibid.; imm. & ; H. and B- 500 mm.; wt. 4-31 kg. 
Macropus (ivallabia) agilis Gould (20) 

(a) Serpentine Creek. Rockhampton district, Queensland; subad. 5 ; H. and B. 
725 mm, wt. 17-73 kg. 

(b) Ibid.; subad. 9 ; H. and B. 643 mm,; wt. 12-27 kg. 
Macropus (ivallabia) dorsalis Gray (21) 

(a*) Mount Lookerbie, Dawson Valley, Queensland; imm. 2 ; H. and B. 
450 mm.; wt. 3-18 kg. 

(b) Ibid.; imm. 6 ; H. and B. 540 mm.; wt. 5-45 kg. 

(c) Ibid.; subad. $ ; H. and B. 570 mm. ; wt. 7-73 kg. 

(d) Spring Creek, Upper Dawson Valley. Queensland; ad. 3 720 mm.; 
wt. 15*90 kg. 


Macropus (-xvallabia) grcyi Gray (22) 

(a) Clay Wells, South-East District, Soufh Australia; aged $ (emaciated); 
H. and B. 648 mm.; wt. 5*97 kg. 

Macropus (zoallabia) parryi Bennett (23) 

(a) Drumburle, Grevillea Plalcam Queensland; ad. & ; H. and B. 805 mil).; 
22 '71 kg. 

(h) Ibid.; ad. 9 ; H. and B. 790 mm.; 15-45 kg. 

(c) Ibid.; imm. ? ; IL and B. 597 mm.; 8*64 leg. 

(cl) Ibid.; imm. $ (large pouch young) ; H. and B. 405 mm. ; 1-82 kg. 

Macropus gigantcus gigantcus Zimmennann (24) 

(a) Coomooboolaroo. Dawson Vallev. Queensland; aged 3 : H. and B. 
1,110 mm.; wt. 50*09 kg. 

(b) Drumburle, Grevillea Plateau, Queensland; subad. 2 : H. and B, 835 mm. ; 
wt. 22-70 kg. 

(c) Yavan Creek, Tumut district, New South Wales : subad & j H, and B. 
950 mm,; wt. 30-90 kg. 

(d) Gorae, West Victoria; imm. $ ; II. and B. 724 mm.; wt. 10*68 kg. 

(e) Jimmy's Creek, Grampian Range, Victoria; subad. £ ; H. and B. 900 mm.; 
wt. 25 '23 kg. 

Macropus gigantcus mclanops Gould 

(a) Padthaway, South-East district. South Australia: imm. 5 ; H. and B. 
576 mm.; wt. 5-45 kg. 

(b) Lake Wangary, Eyre Peninsula, South Australia; imm. £ : H. and % 
850 mm. ; wt. 20-90 kg. 

(c) Coombc, Ninety Mile Plains, South Australia; subad. i ; II. and B. 
870 mm.; wt. 25-90 kg. 

(d) Point Turton, Yorke Peninsula. South Australia; ad. S ; H. and B, 
1,055 mm.; wt. 46-82 kg. 

Macropus gigantcus fuliginosus Desmaresi 

(a) Rocky River, Kangaroo Island, South Australia; ad. 6 ; H and R. 
1,247 mm.; wl. 62*27 kg. 

(b) Ibid.; ad. 5 ; IT. and B. 845 mm. ; wt. 25*22 kg. 
Macropus gigantcus tas-maniensis Le Souef 

(a) Mount Morrison, Ross district, Tasmania: imm. & ; IL and B 775 mm ! 
wt. 18-18 kg. 

(h) Quoin, Ross district, Tasmania; ad. 9 ; H. and B. 835 mm.; wt. 2274 kg. 

(c) Little Tier, North Maequarie River. Tasmania, ad. £ ; H. and B 970 mm * 
wt. 43 '18 kg. 

Macropus rufns Desmarest (25) 

(a) Old Boolcoomatta, North-east of South Australia: aged 6 * H and B 
1,397 mm.; wt. 77*27 kg. 

(b) Tcherricoominyi, Lower Diamantina River (25), South Australia* ad 9 * 
H. and B. 970 mm.; wt 33*18 kg. 

(c) Donald's Well, Musgrave Range, South Australia; imm. S ; H and B 
880 mm.; wt. 20*04 kg. 

(d) Wollara, Basedow R^ng^ area. Central Australia; imm. 2 ; H and B. 
580 mm.; wt. 7-72 kg. 

Macropus robustus robustus Gould (26) 

(a) Coomooboolaroo, Dawson Valley, Queensland; imm. 9 ; H. and B. 
570 mm.; wt. 6'81 kg. 
Macropus robustus ernbr.sccits Sclater 

(a) Mount Norwest, Willouran Hills, South Australia; ad, a ; H. and B. 
1,020 mm.; wt. 50-00 kg. (27). 

(b) Ibid-; ad. 2 ; H. and B. 800 mm.; wt. 2272 kg. 

(c) Ibid.; imm. g ; H. and B. 740 mm.; wt. 15-68 kg. 

(d) Oraparinna, Flinders Range, South Australia; subad. i ; H. and B. 
970 mm.; wt. 37*73 kg. 

Macro pus robustus woodzvardi Thomas 

(a) Deception Creek, James Range, Central Australia; ad. $ ; H. and B. 
890 mm.; 40 -22 kg-* 

(b) Ernabella, Musgrave Range, South Australia; imm. & ; II. and B. 
748 mm. ; wt. 20*00 kg. 

(c) JbuL; imm. 2 ; H. and B. 715 mm.; wt. 15-45 kg. 

(d) Oparinna, Musgrave Range, South Australia; imm. 8 (pouch young); 
II. and B. 355 mm.; wt. 909 g. 


Phnscoloinys mitchelli Oweti 

(a) Hilias Brook, Tumut district, New South Wales; aged 3 ; H. and B. 
960 mm.; wt. 23*18 kg. 

(b) Pecaree, South-East district, South Australia; subad. ■& ; H. and B- 
800 mm.; wt. 19-54 kg, 

(c) Unci; subad. 9 ; H. and B. 850 mm.; wt. 21*81 kg. 

Phascolomys itrsinus Shaw 

(a) Palana, Flinders Island, Bass Strait, Tasmania; ad $ ; IT. and B. 830 mm.; 
wt. 21-36 kg. 

(b) Ibid.; ad. & ; H. and B. 807 mm.; wt. 16*59 kg. 

(c) Ibid,; imm. 2 ; H. and B. 660 mm.; wt. 7*27 kg. 
Phascolomys tasmaniensis Spencer and Kershaw 

(a) Cradle Valley, North-west Tasmania: ad. £ ; H. and B. 935 mm.; 

wt. 27-22 kg' 
(h) Ibid.; ad. 9 ; H. and B. 875 mm.; wt. 22*72 kg. 

(c) Quoin, Ross district, Tasmania; subad. $ ; H. and B. 805 mm.; 
wt. 20-90 kg. 

(d) Meadsfield, Bothwell, Tasmania; imm. 9 ; II. and B. 695 mm. ; wt. 977 kg. 

Lasiorhinus latifrons Owen. 

(a) Portcc, River Murray, South Australia; ad. 5 ; H. and B. 991 mm.; 
wt. 25-45 kg. 

(b) Ibid.; ad. ? ; H. and B. 1,020 mm.; wt. 27-27 kg. 


Pteropus scapulatus Peters 

(a) Coomooboolaroo, Dawson Valley, Queensland; ad. 9 ; H. and B. 230 mm.; 
wt. 280 g. ca. (28). 


(b) Cape Kersaint, Kangaroo Island, South Australia: ad. 8 I IT and B. 
190 mm. ; 270 g, 

Arclocephalus cinereus Peron 

(a) Outer Waldegrave Island, Eyre Peninsula, South Australia: imm g ■ 
II. and B. 1,000 mm. ; wt. 21 36 kg. 

Cams familiaris dingo Blumenbach 

(a) Oparinna, Musgrave Range, South Australia; ad. & ; II. and B. 870 mm.; 
14-09 kg. 

(b) Ooleebinna, Musgrave Range, South Australia; ad. $ ; cranial length 
211 mm.; wt. 16*36 kg. 

Vulpcs vulpes Linne 

(a) Oraparixma, Flinders Range, South Australia; ad, 9 ; IT. and B 550 mm* 
Wt 6-16 kg. 

(b) Dashwood's Gully, Mount Lofty Range, South Australia; aecrl $ ; H and 
B 700 mm.; wt. 5.68 kg, 

Cervus (Dama) dama Linne 

(a) Little Tier, North Macquarie River, Tasmania; imm. $ (feral)- H. and 
B. 1,000 mm.; wt. 19-09 kg. 

Oryctolayus atnicuhts Linne 

(a) Moutajup, Grampian Range area, West Victoria; ad. $ ; H and B 
495 mm.; wt. 1*75 kg, 

(b) Dashwood's Gully, Mount Lofty Range, South Australia; ad. S • H and 
B. 465 mm.; wt. 2- 16 kg. 

(c) Cooncheri, Lower Diamantina River, South Australia; imm. 9 j H. and B 
400 mm. ; wt. 680 g. 

(d) Chundrinna, Everard Range, South Australia; ad, 6 ! H. and B. 425 mm ! 
wt 2-04 ks. 

(c) Spilsby Island, Spencer's Gulf, Soutli Australia; ad. t ; H« and B. 
485 mm.; wt. 1*70 kg. 

Lepus europaeus Linne 

(a) Julia Range r Lower North district, South Australia; ad. ? ■ II and B 
590 mm.; wt. 3-07 kg. 

(b) Point Turton, Yorke Peninsula, South Australia; ad. 2 ; H and B 
615 mm.; wt. 3- IS kg. 

Rattus z'iHosissimus viUosissinnts (Waite) (29) 

(a) Appamunna, Lower Diamantina River, South Australia; ad. £ ■ H and 
B. 188 mm.; wt. 185 g. • ' 

(b) Ibid,; imm. 8 ; H. and B, 140 mm.; wt. 90 g. 

(c) Ibid.; ad. 5 ; K. and B. 195 mm.; wt. 175 e> 
Ratius greyi greyi Gray 

(a) Callawonga Creek, Fleurieu Peninsula. South Australia; ad <5 \ H and 
B. 139 mm.; wt. 82 g. 

(b) Boat Harbour Creek, Fleurieu Peninsula, South Australia: ad 5 • H and 
B. 157 mm.; wt. 112 g. 

(c) Ibid.; ad. ? ; IT. and B. 139 mm.; wt. 90 g. 

(d) Ibid.; imm. 9 ; H. and B. 122 mm.; wt. 51 g. 


Ratfus greyi peccahts (Troughton) 

(a) Heathmero, West Victor/a; ad. 2 ; H. and B, 159 mm,; wt. 100 g. 

(b) Ibid.; ad, <$ ; H. and B. 169 mm.; wt 130 g. 

Rattus luircola httreola Gray (30) 

(a) Rat Island, Lake Alcxanririna, South Australia; ad. 6 ; H. and B. 
173 mm.; wt. 190 g. 

(b) Ibid,; ad. 2 ; H. and B. 178 mm.; wt. 190 g. 

(c) Wannon River. Yarram Gap. Grampian Range, Victoria; subad. 2 ; 
162 mm.; wt, 148 g. 

(d) Blacktellows Creek, Flcurieu Peninsula, South Australia; subad. £ ; 
H. and B. 156 mm.; wt. 130 g. 

(e) ITeathmere, West Victoria; imm, 5 ; H, and B. 134 mm.; wt. 70 g. 

(f) Mount Clay Range, West Victoria; imm. 9 ; H. and B. 98 mm.; wt. 26 g. 

Rattus rattus rattus Linne. 

(a) Mackenzie River, Grampian RaUgK. Victoria; ad. 9 ; H. and B. 179 nun.; 
wt. 168 g. 

Rattus rattus alcxaudrinus Geoffroy 

(a) Jimmy's Creek, Grampian Range, Victoria; ad. tj ; IT. and B. 189 nun.; 
wt. 178 g. 

(b) A garden, Adelaide, South Australia; a<l 2 ; H. and E. 190 mm.; 
' wt. 190 g. 

Ratfus norvegicus Rrxleben 

(a) Mcadsneld, Bothwell district, Tasmania, ad. 3 ; II. and B. 262 mm.; 
wt. 445 g. 

(b) A garden. Adelaide, South Australia; subad. 2 ; II. and B. 206 mm.; 
wt, "200 g. 

(c) Ibid.; Imm. d ; II. aud B. 115 rnm,; wt, 100 g- 

Mas muscuhts Linne (31) 

(a) A house, Adelaide, South Australia; ad. # ; H. and B. 83 mrn.; wt. 19 g. 

(b) Ibid.; ad. 2 ; H, and B. 86 mm.; wt. 22 g. 

(c) Boat Harbour Creek, Fleuricu Peninsula, South Australia; o ; II. and B. 
83 mm.; wt. 11*5 g. 

(d) Ernabella, Musgrave Range, South Australia; ad. o ; H. and B. 80 mm.; 
wt. 9 g. 

Pscudomys (Pseudomys) higijinsi Trouessart (32) 

(a) Cradle Valley, Morth-west Tasmania; ad. <5 ; II. and B. 132 mm.; 
wt. 75 g. (approxj. 

Pscudomys (Pscudomys) minnie Troughlon (33) 

(a) Appamunna, Lower Diamantina River, South Australia ; aged <5 ; II. and 
B. 131 mrn.; wt. 80 g. 

(b) Ibid,; ad. 2 ; H. and B. 134 mm. ; wt. 75 g. 

(c) Ibid,; imm. o ; H. and fk 115 mm.; wt. 44 g. 

Pscudomys (Thetomys) nanus Gould (34) 

(a) Koonapandi, Musgrave Range,, South Australia; ad. 6 ; H. and B. 
101 mm.; wt. 28" 5 g. 


Pseudomys (Laggadwa) hcrmannshurgensis hcrmonnsburgcnsis Waite (35) 

(a) Wollara, Basedow Range area, Central Australia; ad. o ; H. and B. 
74 mm.; wt. 14*5 g. 

(b) Ernabella, Musgravc Range, South Australia; ad. 9 ; H. and B. 80 nini; 
wt. 13 £. 

(c) Chundrinna, Everard Range area. South Australia; subad. 9 ; H. and B. 
71 mm. wt. 7'5 g. 

Pseudomys (Gyamys) apodcmohlcs Finlayson (36*37) 

(a) Coombc, Ninety Mile Plains, South Australia; ad. g ; H. and B. 86 mm.; 
wt. 16 g. 

(b) Ibid.; ad. 9 ; H. and B. 93 mm.; wt. 18 g. 

(c) Ibid.; imm. o ; H. and B. 70 mm.; wt. 9 g. 

Mastacomys fuscus Thomas (38) 

(a) Cradle Valley, North-west Tasmania; ad. 9 ; H. and B, 168 mm.: 
Avt. 160 g. (approx.). 

Hydromys chrysogasler f ul vol ava tits Gould 

(a) Fulham, Torrcns Creek, South Australia; ad. 6 ; H. and B. 343 mm,; 
wt. 985 g. 

(b) Ibid.\ ad. 9 ; H. and B. 310 mm.; wt. 700 g. 

(c) Mosquito Creek, South-East district, South Australia; imm. <S ; H, and B. 
292 mm.; wt. 390 g. 

(d) Furnen South-East district, South Australia; imm. 9 ; H. and B. 293 mm.; 
wt. 425 g. 

IlyxlrmnXA ihfyMffimtff "mf, (Sir) 

(a) Innamineka, Barcoo River, South Australia; ad. 9 ; H. and B., 355 mm.; 
wt. 500 g. 

The references are, in all cases, to papers by FT H. Finlayson, published 


Trans. Roy. Soc. S.A., as follows: 


1933 ; 

; 57, 197 





1933 ! 



1935 . 



1932 ! 

i 56, 168 


1935 : 


1934 : 



1930 : 

54, 177 


1934 : 

58, 224 


1934 : 

58, 220 


1934 : 



1935 : 

59, 223 


1935 : 


27. 1936 : 60, 158 

28. 1934 : 58,230 

29. 1939 : 63, (I), 88 

30. 1935 : 59, 224 

31. 1939 : 63,(1), 115 

32. 1933 : 57, 129 

33. 1939 : 63, (1),94 

21. 1931 ; 55,73 34. 1941 \ 65, (2), 224 

22. 1927 : 51,363 35. 1941 : 65, (2), 215 

23. 1931 : 55,75 36. 1932 : 56,170 

24. 1931 : 55,68 37. 1944 : 68, (2), 210 

25. 1936 : 60, 157 38. 1933 : 57,125 

26. 1931 : 55,69 39. 1939 : 63, (1), 114 


1932 : 

; 56, 148 


1936 : 



1931 ! 

55, 85 


1931 . 



1930 ; 



1930 : 



1931 : 




By Allan F. Wilson 


This paper is the first of a proposed series dealing with the geology of the Musgrave Ranges. Some 
features of the major rock types are outlined below, and other papers are in preparation which will 
amplify and supplement this introductory account. Ultimately the petrogenesis of this new series of 
"Charnockitic" rocks will be treated. 




Fly Allan F.. Wilson* 

[Read 8 May 1947] 

Plates II to IV and Geological Map 


a. ^troductiux 

PrevJous Work 1<)5 

Location and Terrain . . . . . . . . . . . . . , . . . . - . 196 

Locality Plan r , 199 

Concerning maps and methods .. .. .. . . .. .. .. .. 196 

The term "Charnoclcitc" . . . . , , 196 

B. Major 'Rock Types 
I. The Gneisshs 

Umbyarra to Kenmore Park , . ., .. , * .. .. .. . . 198 

The major area — general features of the gneisses . . . . .. .- . . K*9 

(1) Gneisses showing Scd : mcrtary origin . . , . . , . . , , . . 200 

(2) Gneisses not showing Sed'menlary origin and containing no 
fluorescing zircon . . ► . . . . . - , . . . ~ - , . ., 201 

(3) Gneisses not showing Sedimentary origin and containing 
fluorescing zircon • • 203 

II. The Non-Gneisstc Ch^rxockttes 

(1) The Mt. Woodrofre and Trudingtr Pass Intermediate and Basic 

Rocks 204 

(2) The Upsan Downs Gran'-tic Rocks .. .. n fi .. .. 206 

( 3) The Norite to the S.W. of Tjatjanja . . . . . - ■ 206" 

(4) The Ernabella Granodiuritic Mussif 200 

III. Pegmatites .. .* .. 208 

IV. Dolcritcs , . . , . 208 

V. My Ionization 209 

C. CONCt/USlOX AND SUMMARY' .. .. 209 

D. Acknowledgments . . . . . . . . . . . . . . . . - . . ., 210 

E. Bibliography . . . . . . . . , . . . . . . , . . . . ... 21 


This paper is the first of a proposed series dealing with the geology of the 
Musgrave Ranges. Some features of the major rock types are outlined below, 
and other papers are in preparation which will amplify and supplement this intro- 
rluctory account. Ultimately the pedogenesis of this new series of K Clia£ft0Ckiticr 
rocks will be treated. 

Previous Work 
A bibliography of previous workers in the Musgrave Ranges is appended. 
The two most notable contributions to our knowledge are those of Basedow (1905 ) 
and Jack (1915). Basedow's account covers the reconnaissance of the South 

* Geology Department, University of Adelaide. 
Trans. Kay. Soc. S. Aust., 71, (2), 1 December 1947 


Australian Government North-West Prospecting Expedition of I 903 which 
penetrated farther west than the Mttsgraves, into the Mann and Tomkinson 
Ranges and to the "Western Australian border, jack's work was carried out in &9 
abnormally dry season (1914). Consequently most of his time was spent to the 
south-east of the author's area. Notwithstanding the account? of both Basedow 
and Jack contain valuable observations, to which reference will be made in later 

Locality avp Ti-iutAiN" 
The Musgrave Ranges are the easternmost of a series of rugged ea-i-wcst 
.mountain chains in the tar North-West of South Australia. 

These ranges begin in the east with hold desen-red Sentinel Hill (180 mil--; 
west of the Adelaide to Alice Springs railway line at Finke, Northern Territory), 
| See locality plan, p. 199.] As one continues westward from SenLinel Hill the 
Mu^t-avcs groW in magnitude and area, but are never more than 30 miles from 
north to south. Tbey 'reach their culmination in the mountains in the vicinity 
of Mount WoodroiTe" 5.000 Eetft above sea level, and 3.000 feet above the desert 
plain. This range continues west for over 1C0 miles, but a spur parses west- through Opparitwa and thence into the Petcrmann Ranges within the 
Northern Territory. To the west of the Musgravcs, the Mann and d:en the 
Tumkinson Ranges continue almost into Western Australia. 

A characteristic feature of the Musgrave Ranges is the abrupt manner in 
nhich most of the mountains rise from the plain. Groups of precipitous island-like 
hills punctuate the flat plains, which are bounded on either sirje by precipitous 
mountain ranges. Such ruggfed scenery creates a strong impression of somr 
Norwegian fiord. This illusion is greatly accentuated by the lengthening shadows 
of evening. 

The maia area under consideration (see map) comprises the ranges from 
Kcnmore Park (approximately longitude 132° 30' K.}, westward for 50 niiles^to 
Mourn WoodrottV and Ertywafiyawanya (approximately longitude LIT" 40'). 
Latitude 2(> 3 15' S. runs through the centre of. the ranges. Most detailed work, 
however, was done within a few miles cd" the Presbyterian Mission Station at 

The period of the two field operations undertaken in the Mitsgravcs wa-s 
during due mttei exhausting time of the year — that of the Central Australian 
sumrricr. The first occasion was during December 1943, January and February 
1944, followed bv a second in December 1945 and January 1946, these periods 
being the only times available, tor the purpose. With Krnabel'a as the base oi 
the survey, the ranges were investigated by horse, camel, and motor truck-, but. 
large! v on fool. Transport difficulties and seasonal conditions limited the most 
intensive work to a radius of 10 mile.-- from the base. Considerable reconnaissance 
and some detailed work was carried out over a wide area, as far west as Mount 
WoodrofTe and Erlywanyawanya, and Kenmore Park m the east. Positions uf 
enmp sites and consequent geological work were often hampered by water 

Concerning Mats and Methods 
in 1892 Carruthers published a map of the north-west of South Australia. 
on a scale of & inch to 1 mile. This appears to have been the basis of maps used 
by all later expeditions to the north-west. When a map of the Central and 
Eastern Musgraves was needed, existing 1 maps, which had their uses for general 
purposes, were inadequate for more detailed work. Hence T was compelled to 
make my own maps, and used rough theodolite and compass methods for speed. 
The nix-tri.£g-ed point? of Carruthers which fell withm my area were incorporated. 


Willi the exception of that of Mount Everard. The actual location pi flic lartor 
is unknown to the author and to all at Ernabella. Nor do the natives know ot 
any triced mountain in the plotted portion of Mount Everard. The map of the 
vicinity of Ernabella was prepared on a scale of 4 inches la 1 mile. The larger 
area was prepared on a ^cale of J inch (o 1 mile, tlftylS Kgni£ of Carruihcr^' data 
when tny own dnia and lime were insufheiem. Numerous panoramic, photographs 
were used io check as much detail as po-sible. Some truck mileage-: n-n. 

Owing to difficulty in location of parts of the track from Ernabella to Ken- 
more Park, reliance had to be placed on truck mileages and compass readings. 
With the truck "mileages*' m need of calibration certain parts must be regarded as 
sketch maps. This applies also to the area jnst north of Alalka, the vicmiiy of 
tbe "pass** to the east of Ombaguuda. near Fig Tree Gully Soak and JJjsSMl 

A prismatic compass ha? only limited use in these ranges. The magnetic 
varialiou was found to be 4-V 3 to the east of true riorlh. The rocks of the area 
greatly afTecl the compass, so much so that great care must be taken ii that 
instrument is to be u=ed among the mountains. When on the plains, accurate 
readings can be obtained. The magnetic variation is usually very erratic. Even 
nn the same hill I have noticed a difference of up to 5° to 6° between readings 
taken dining and .standing although exactly in the same place, Uearin^s taken 
when one is enclosed in rugged ravines are often well-nigh useless. Many hear- 
ings were, found, on checking-, In be hopelessly inaccurate. The only way to take 
<i reasonable compass reading* was found in be by standing in such a position that 
t!ie interfering rock is equally distributed on all sides. Such being tbt cast-, a 
compass survey was often found most i^c.ious. Some type of sun-compass would 
<'iuibtless be easier to manage. 

Aboriginal names were used on the map where possible. Care must be taken 
ivith these; however, for native place-names are rardy as precise m meaning as 
our own. For instance, while pointing to Mount Spec, the author was told that 
th aboriginal name is "hiindi." But it was later discovered that the mountain 
lias no particular native name, but that "Jnindi" refers to the "region" of the wck 
hole situated at a considerable distance to the south-west. The probable explana- 
tion Is that to the aboriginal the tormidable and barren Mount Spec has no 
need for a name, so when pestered by the white man for one, the nearest he can 
give is ' luindi.''* Similarly, £ *Tjatjanja u (or fi Tjatja") probably refers to a rock 
hole the south of the mountain gjvtjfl that name. 'Ttjinpirt" io the aboriginal 
denotes the series of rock holes to the east of the imposing mountain mass which 
(he missionaries at Ernabella call Mount Itjinpiri. 

In the absence of suitable aboriginal names several were coined. The origin 
of of these will be obvious, bur some need explanation. "Trudinger Pass" 
(suitable for horses but found difficult for camels) was named after Ronald 
Trudingcr, friend and host at the Ernabella Mission. "Kroek Pass" (probably 
negotiable with camels) was named after Richard Brock, who accompanied the 
author on the second expedition, "Henderson Rock Hole" is a fine water about 
which the aboriginals apparently know very little because i{ is just above a big 
cliff and away from normal hunting grounds. It was discovered while descending 
Mount WoodrofYe. and was named after Ronald Henderson, who Wilfully drove 
l he truck through the nearby difficult Brown's Pass. 

The large bare water catchments of the Musgrave Ranges make rock holes- 
fairly common, but very Few are reliable. On the accompanying map all reason- 
ably useful waters arc shown, that is, I hose which were found with some water 
during the two dry seasons of the authors visit. 


When vertical aerial photographs are taken (2 > many parts of the accompany- 
ing map will, uf course, need to be somewhat modified. However, it i? considered 
that the map is a definite contribution to our present knowledge of the area, and 
little difficulty should be experienced in locating any position marked on the map. 
Therefore, apart from the geology represented, the location of various features 
of the area (rock holes, passes, etc. ) fcems to justify its publication. 

The Tkhm *'CuAj?\-<>eKrrr." 

The ranges comprise a complex mixture of gneisses and ancient bul later 
deep-seated non-«nth*ssic hrneous rocks. Numerous dolerire dykes cul all of these 
earlier systems. All rocks ot the Musgrave Ranges are of Precambrian age 
excepting, of course, the thick mantle of Recent red desert sands and loams of 
the valleys between the ranges. 

Most of the Musgrave Ranges rocks, both ;he gneissic and non-gneissic types, 
are hypersthene-bcarin^. Notable papers have been published in i _ eceut yea,r.s 
treating Charnockites froui several regions of the world, and it appears from the 
divert origins which are claimed for these hypersthenc-bcaring igneous rocks, 
gneisses and granulites, that the term "Charnoekite" has lost much of 
what it was originally intended to convey. However, for the present. 
until petrogeuetic relationships in the Musgrave Ranges have been more fully 
interpreted, these rocks may well be called "The Charnockites and associated rocks 
mi North-Western South Australia." 

I. The Gneisses 
l"\rnv\RKA to Kexmore Pakk 

At Umbyarra Bore ancient gneisses were first noticed on the track ue^t from 
Kinke. Umbyarra is 64 miles west of Finke in the Nortberu Territory. They 
outcrop as low hills amid remnants oi* horizontal sandstones of the late Meso/x»ic 
and early Cainozoic. At Umbyarra some pegmatites were observed, but most 
of these arc barren. One pegmatite contains gaod crystals of muscuvite and some 
tourmaline. The general lrend of the pegmatites is N25~ R. To the west the 
<iTie ; sses are more extensive, though they rarely form any more than low hills and 
inconspicuous outcrops in the mul^a scrub. 

About 2 miles west of Umbyarra Bore a prominent hut low range of hills, 
about a mile and a half to the north of the track, was visited. This is probably 
typical of similar low lines of hills which protrude at intervals from the surround- 
ing flat country. The hills consist of a non-foliated biotite-beariug granite; the 
flanks of the bills show excellent contacts with the ancient gneisses. Numerous 
chunk:* of gneiss were noticed to be partly "assimilated" by the granite. A few 
barren pegmatites are present. 

Similar granitic gneisses continue to the west, but become more heavily 
injected b> east-west dolerire dykes towards "The (lap/* 122 mi!es from Finke. 
and approximately on ihe South Australian-Northern Terr'lory border. At '"The 
Gap'"' the gneisses show considerable crush. The dolerites are similar n» those of 
the Musgrave Ranges. Basedow made observations concerning parts of this area 
of the "Avers Ranges"' ( Masodow 1905, p. 78). 

P? Within :i few day^ of pre.senttnft tin* paper to the Society a batch of 27 tow- 
level oblique aerial photograph? were sent to t!ie author by courtesy of the Australian 
Geographical Society, were uiK-en with a view to publication in "Walkahout," 
lie Org^n of the society, hence the prime purpose of the photographs \\'a=. not carto- 
graphic. They show the northern part of the Ranges near Mount Woodroffc. Trudinger 
Pa.5S Alallca, Tcdj.ii'itja and Wed^e Hul. The photographs reasonably confirmed the 
acrnracy of the mop in these regions. A few minor alterations were made. 


At Beefwood Creek, 135 miles west of Finke and a few miles south of the 
Northern Territory border, an interesting pyroxene granite intrudes the acid 

From Beefwood Creek the types of gneiss vary more frequently. The 
most common is granitic, but intermediate and basic types are plentiful. At Ken- 
more Park, nearly 180 miles west of Finke and just within the area covered by 
the map, the variety of gneisses is great. Garnet-bearing and/or hypersthene- 





Fig. 1 Locality Plan 

bearing gneisses are abundant. The general trend of all of these gneisses from 
CJmbyarra to Kenmore Park is north to south. The dip varies from 6Q°-70° to 
the east or west. 

Ttte Major Area and General Features of the Gneisses 
The area to the west of Kenmore Park was studied in more detail. From 
the map, the importance in this area of gneisses and granulites is readily seen. 
Most of these gneissic rocks are a dove-grey colour with streaks of dull dark 
brown pyroxene and occasional black shining amphibole. 


The map shows that the gneisses commonly have a north south strike w with 
a steep dip to the cast. Towards the north-western part ol the area, however, 
the strike of the gneisses swings around from a few degrees west of north to 
take on an almost east-north-east trend near Knna-unpunj'a and Warclulka, The 
dips are such as to suggest that the western half of the urea is part of the western 
iimb of ;i south-pitching syndine. Anomalous dips and strikes arc not uncommon, 
hut most of these are close to junctions with the non-gneissic igneous rocks. u> 
T&feac may represent minor buckling, reversal of dip and possible ovcrthrustini? 
related to the origin of the igneous rocks. Further detailed work is needed in 
critical areas. 

The question of the origin of these gneissic rocks is difficult, and h would 
be premature at this stage of the investigation of the rocks of this region to state 
final conclusions as to their pedogenesis. However, certain suggestions are put 
forward, and these are linked with the ideas of "palingenesis" and "granitiza- 
tiori'* considered as having acted on a grand scale (see p. 209). 

From the held evidence the author considers that the area consists of a 
series of ancient highly metamorphosed sediments whose eventual granirization 
has brought about most of the non-gneissic igneous rocks. However, in hand 
specimen, the sedimentary origin of the gneisses is usnalry not so obvious. 

The following- observations on some of ihese gneisses are not primarily con- 
cerned with the setting oxtt of petrogenetic relations. These will appear in sub- 
sequent papers. 

However, the order of the gneissic types is probably chronological; thai is, 
the rocks first described are the oldest. 

(1) Gneisses showing- sedimentary origin. 

(2) Gneisses not showing; obvious sedimentary origin, 
(o) Acidic gneisses without fluorescing zircon, 

(b) More basic gneisses without fluorescing zircon. 

(3) Gneisses not showing obvious sedimentary origin, hut showing 
fluorescing zircon. 

(1) (Jxeissks showing Sedimentary Origin". 

The gneisses showing nn obviotts sedimentary origin are not plentiful. The 
recognition of rocks as of sedimentary origin can often be difficult. In this area of 
chamockitic rocks, plutonic conditions resulting* in palingenesis and granitization 
are assumed to have been in force for considerable periods. Rocks of the arkose 
type, if present under such conditions, could be presented m such a form as to 
defy recognition of their origin. Argillaceous and highly calcareous rucks probably 
could be more easily identified. 

No rock was found which obviously came trom a highly calcareous rock. 
but there are some for which an argillaceous origin is -suspected. These areas 
are of limited extent : 

(1) part of the complex gneissic system to the. south-west of Mount Car- 
ruthcrs ; 

p5 The term "gneiss" is here used for rocks showing varying degrees of batuhnu. 
The ward "gneiss'* is preferred to "foliated rode," since "the banding is considered to 
represent the bedding- planes of the original sediments, now gneissificd. The terms "dip" 
and "strike" are therefore used (o describe this banding. No schists were noted. 

t-> The term "iiou-gneissk igneous rock" is meant to convey the idea that u\t 
major igneous rocks of this area have been derived from sediment^ the gneissic struc- 
ture of which has become obliterated in the process of formation of the "igneous" UiaSSeS. 


(2) possibly the greater part of the gneisses to the north-west of Palpatjara 
Weil ; 

(3) small patches 1 mile to the imnh-wefct of Taljariijn and lh mile-, to the 
west-suiith-wCi-t of Taljaritja; 

(4) patches 1 mile north-west of Top Springs; 

(5) patches up the Krnahella Creek from Top Springs; 

(6) some of the gneisses to the north-east of Eriywanyawanya B-oek Hole; 

(7) 2£ miles due west of Mount Camtthcrs; 

(8) 1 mile north-east of Taljaritja Well. 

These rocks contain variable amounts of at least four of the following 
minerals". — game', cordierite, sdlimanitc, biorite and a green spinel. The garnet 
is always visible in hand specimen r.s pink rounded crystals, usually about 2 mm. 
in diameter. The cordierite in the held is noi noticeable in most specimens, though 
the presence of a "greasy" dull yellowish-brown with the garnet leads one to 
suspect that mineral. (Hut v 'greasmess" is by no means necessarily indicative of 
cordierite, for most of the hypersthene.-hearing rocks of this area show some 
"greasiness"..) Only in one locality, (7), lias macroscopic sillimanite been 
observed. It occurs as brown crystals up to 2 cm. long. The dull dark green 
spinel shows very high ahsorpiion. so much so that with a cursory glance at a. 
slide, one could quite easily overlook if, grouping it with the iron ores. It usually 
occurs intergrown with iron ores, SitHmanke is commonly aligned within the 
spinel,, but ar localities (7) and (8) the siilimanite also occurs as plentiful 
euhedral crystals. In all of these rocks bioti'.c is plentiful, However, in the 
normal gneisses and non-gneissic charnockites to be mentioned below, hiotitc 
occurs usually as a very minor accessory. 

It should be stated that the presence of garnet in acidic rocks must not he 
taken as necessarily indicative of direct sedimentary origin in this region. Garnet 
has been noted in many other places, but usually near the junctions of the main 
Rruabelia non-gneissic "igneous" mass and in the gneisses around certain 
"cupolas" to the east (e.g., Spinifex Hill), and often in mylonized zones. 

Some analytical data and petrographic descriptions have been assembled, but 
will be withheld for inclusion in later papers. However, it may be stated that in the 
analysis of an ''injection gneiss 1 ' over 5% corundum is indicated in the norm. 

The author considers that the rocks which show evidence of sedimentary 
origin are often greywackes ot perhaps relatively high alumina content, and that 
later investigation may show that large tracts of the gneisses can claim palin- 
genctic relationships with arkosic rocks. 


(a) Acidic Gneisses zc'ittwitt l 7 !icurvsiittf/ Zircon. 

Many specimens representing over SCO localities have been carefully studied 
under ultra-violet light. In large numbers, a zircon has been revealed us tiny- 
points of orange light. Sometimes the crystals arc up to 5 mm,, but usuallv of 
the order of 1 mm. in length. In most of the gneisses, however, no fluorescence 
was observed, and petrographic work has revealed that a somewhat different non- 
ftuoreseing zircon (?) exists in these rocks. The author hopes that it may be 
possible lo trace petrogenKic relationships by making use of this zircon criterion. 
The areas represented by the fluorescing zircons are indirated in the sketch map. 
p. 202. in hand specimen, little indication is given of the presence or otherwise 
of the fluorescing zircon. However, those with fluorescing zircon do .seem to 
exhibit a somewhat more "igneous" appearance. 


The present indications are, however, that the gneisses without the fluorescing 
zircon are somewhat older than those with the fluorescing zircon. Investigation 
into this interesting phenomenon is at present only in the early stages. 

A number of modes have been established by the Rosiwal method, and some 
analytical work has been undertaken. (Details will appear later.) From this 
it was found that all gradations from granitic to granodioritic rocks occur within 
this acid group. An average type (adamellitic) occurs at the little rock hole, 
2 miles west of Top Springs. 

This group of gneissic and sometimes granulitic rocks are dove-grey and 
of a greasy lustre. The dark minerals are usually concentrated into rough, 
more or less continuous plates throughout the rock, and there arc usually some 
rude bands of feldspathic material. These strike north-south, with a fairly steep 
dip which varies from east to west. The rocks are usually fine-medium grained. 
Microscopically these gneisses and granulites usually show rather rounded yet 

Fig. 2 

interlocking grains, with the ferromagnesians of slightly smaller dimensions than 
the salics. Some quarts grains are particularly ragged and associated with 
myrmektte, while others protrude as small irregular ''angers" through the rock. 

The plagioclases are usually twinned on albite and pericline laws with 
composition about andesine (An. 35). They are variable in amount, but are 
commonly plentiful. Myrmckite is often associated. Quartz at times contains 
numerous non-orientated needles. Shadowy extinction is common. The "fingers 1 ' 
of quartz are mentioned above. 

The potash feldspar is usually abundant and represented by an orthoclase 
mieroperthite, but sometimes by a poorly twinned microcltne rnieroperthite. The 
proportion of the other minerals varies greatly, but hypersthenic is usually the 
commonest. The pleochroism is strong in rusty pinks and greenish-greys, with 
a slightly variable optic axial angle (negative) of about 80°. This pyroxene 
often shows considerable cracks filled with greenish alteration products. A pa!c 
green non-pleochroic monocline pyroxene (probably diopside) is usually present, 
but hypersthene predominates. 


Hornblende is sometimes relatively plentiful blit is often absent. Pleochroism 
is usually yellowish-grey, light brown, dirty green, with Z Ac 23 J -25 Q . 

Magnetite is often abundant and at times an important constituent. 
Secondary magnetite is commonly developed near hypersthene. P.iotite is asso- 
ciated iti'smali amount with the iron ores, Pleochroism is pale straw-yellow to 

Zircons ( ?) : present indications are that all of these crystals, which are non- 
fluorescent, are usually murky and tending to be. brownish and much cracked- 
Apatite is commonly present. 

On chemical analysis, a typical gnei-sic rock revealed the normal features of 
an "iMieQUs" rock of adame.llitic composition Such rocks couid well be called 
acidic gneissic charnockites. 

(h) Intermediate and -more Basic Gneisses without fluorescing Zircon. 

These are best represented by some of the dark gneisses at Gilpin's Well. 
although patches of similar rocks occur near Kuna-unpunja, Brown's Pass, and 

In these rocks the gneissic structure is usually marked and is revealed on the 
weathered surface by streaks of tiny greyish, ragged quartz crystals, and shiny 
magnetite. On fresh surfaces, specimens arc a bluish dark grey (contrast the 
"dove grey" of acidic gneisses) with the typical greasy lustre of the other 
hypcrsthene-hearing types. 

Microscopically these rocks are granoblastie, with an average grain 
size of the plagioclase just under 1 mm. in diameter, and that of the ferro- 
magnesians about O 5 mm. Modal proportions arc 65%, Hypersthene 
14%, Diopside, 7%, Quartz under 5 c /o, and Hornblende, etc., 9 c /o (Gilpin's Well). 

The plagiodase is usually a well-twinned labradoriuc antipcrthite (An. 45) 
with plentiful opaque needle inclusions. The hypersthene is fresh and displays 
the usual strong pleochroism. 

The monodinic pyroxene is pale bluish green-grey, unn-pkochroic, and has 
Z/\c = about 45°. 

Quartz usually occurs as the finger-like protuberances so commonly seen in 
the more acidic gneisses. The mineral often has needle inclusions similar to 
those in the plagioclase. 

Magnetite is usually quite plentiful in these rocks. The zircon (?) is 11011- 
fluorescing and of the murky type, noted above in the acidic gneisses. Some. 
apatite and biotite are commonly present and associated with the magnetite. 
Potash feldspar and hornblende are very uncommon. 

Such rocks could well be called quajtz-dioritic gneissic charnockites. 

(,3) Gniusses showtxc no obvious Skdimkntar^ Origin a\d contattc- 

iKO Fluorescing 7arcox. 

From the sketch map on p. 202 it can be seen that these gneissic rocks appear 
1o occupy mainly: — 

(a) an east-west belt from soiuh of Mount Wnndroffc through south 
Trudinger Pass and Brock Pass to the Tnindi area; 

(b) the east-west Mount Cuthbcrt Range to the north; 

(c) the east-west belt from Mount Ferdinand to Omhagtmda; and 

(d) many other minor localities. 

The strike of these gneisses is still the same, north to south of the whole region, 
with the dip exceeding 45 * east or we s ;t, excepting in the area near the Wardulka 
Rock Hole [Locality (b) ]. Here the gneisses take on a more of an east-west 


As was suggested above (p t 201), ft seems that these gneissic rocks are a 
little coarser (though still far from coarse-grained) and look more "igneous" than 
the gneisses, free from the fluorescing zircon. 

As in the first group of gneisses, this group shows gradations from rhe highly 
acid to the granodioritic and quartz-dioriric types. The normal grain size is found 
to be about 1 mm, in diameter, with the ferromagnesians perhaps a little smaller. 

The rocks show the customary greasincss of ail the hypersthene-bearing rocks 
of this region. In most, the typical dove-grey colour is evident, though some 
specimens show more of a fawn-grey, owing, perhaps, to alteration, 

Orthoclase microperthite is usually plentiful, but tnicrocline micropcrthite 
is only occasionally present. At times, an acid andesme is the dominant feldspar. 
This plagioclase is commonly autipcrthitic. A quick method for distinguishing 
the perthitic from autipcrthitic feldspar was noted. This consists in "viewing 
rhem under considerably reduced illumination. The amiperthitic particles then 
appeared pale pink, while those of the perthite remained colourless. 

Quartz occurs as ragged tongues" and scattered like millet-seed throughout 
the potash feldspar in various rocks. Occasionally non-orientated needle- 
inclusions are developed in the mineral. 

'fhe dominant ferromagnesian minerals are a typical pleochroie hypersthene 
and a pale bluish-green non-pleochroic diopsnie. These arc commonly arranged in 
rough bands which are separated by the salic-rich bands. Common hornblende has 
been noticed, but it is not typical of these gneisses. Iron ores are often plentiful 
<xnd arc commonly inrerlocked with the pytmeues. 

Apatite occurs in variable amount, but mostly associated with the pyroxenes 
and iron ores, Several crystals of zircon of the fluorescent type occur in all 
slides. The fluorescing zircon is clear and colourless under convergent light, 
whereas the non-fluorescing zircons (?) are almost always murkier and, under con- 
vergent hght, usually show a very pale pinkish-fawn tint, 

ILL The Non"-G;nei5»sic Charxockitks 

Large areas of non-gnetssic rocks <:,) are shown on the map. Most of these 
would usually be called igneous rocks. Though these arc. younger than the 
gneissic racks, the final period of crystallisation of the uon-gneissic "igneous" 
rocks is probably not far removed from that of some of the gneissic types. 

At present these non-gneisstc rocks may he divided into four divisions, in 
this scheme the oldest types are listed first The four divisions are: — 

(1) The Mount WoodrofTe and Trudinger Pass Intermediate and Haste 

(2) The Upsan Downs Granitic Rocks. 

(3) The Norite to the south-west of Tjatjunja. 

(4) The Ernabella Granodioritic Massif : 

( I ) The Mount Woodroffe - Trudiaxi£k Pass I .vrruMEDiATE akp Ba^ic 

(a) The Mount Woodroffc Area, 

The. map indicates that on ail sides of the Mount gneissic rocks predominate, 
but near the summit considerable areas of cjuartz-diorilic and dioritic rocks occur. 
There are basic and ultrabasic patches in places. Directional characters were not 
noted in any of these charuockiue rocks on Mount Woodroffc. However, the 
strike and dip of the surrounding gneisses, and the comparison with the basic 

t s " Sec (ootoeie W on j>. 2CH>, 


rocks which continue iu a north-easterly direction to Tni dinger Pass, suggest that 
these rocks prohahly comprise a large sill-like structure. This is more evident in 
the range between Brown's Pass and Landmark Hill. Directional characters 
were not noted in any of these rocks, bur some may be expected when field work 
can be carried out on the cross-section of the apparent sill structure. <G) Recon- 
naisauce only has been possible in this area up to the present. 

in hand-specimen the rocks of Mount WoodrofTe are medium-drained, con- 
sisting of pale amethyst grey and greasy kit shiny plagioclase, mottled with 
patches of dull black pyroxene. 

Twin striae are not easily seen in the plasrioclase. in hand-specimen, hut under 
the microscope albite-rwins are plentiful. This antiperfliitic feldspar (70%) is 
a rmd-andesine. 

Hypersthenc usually occurs a? large tabular rounded crystals about 4 mm. x 
3 mm r It is very strongly pleochroic. Diallage is also present. Quartz h noted 
in accessory amount. Potash feldspar has not yet been seen except as the anti- 
perthitic lenses. Zircon is quite rare and of the nori-fluoreseing type. 

Hypi'-rstlienites appear as patches up to an acre in area on the sides of Mount 
Woodroffe. Another outcrop occurs in Brown's Pass between the Woodroffe 
tocks and the basic rocks stretching north-east to Trudinger Pass, These ultra- 
basic rocks are commonly associated with magnetite and very coarse dtnritie. and 
nuritic rocks similar in appearance to those of the summit described above. The 
hvpersthenites are fresh, dense homoRciieous medium-grained, greasy and very 
dark grey rocks. Microscopically, they are found to consist almost entirely of 
pvroxene", most of which is a strongly pleochroic hypersthenc. There is a little 
non-plcochroic pale green dialla^c. Plagmclase is usually present in extremely 
small amount, as tiny interstitial particles, Very little primary iron is present in 
slides studied to date, but in the field, solid boulders of magnetite arc common 
as floaters near the hyperslhenires. Directional features were not observed. 

(b) The Trudiiujcr Pom Btuic Itocfcs. 

Ak suggested above, these wvritit* rocks are linked with the Mount Wood- 
roffe rocks, ' The outcrops m the Pass were noted during a particularly arduous 
rcconnaisauce on camel, and time could not be spent trying to determine the 
relation of these rocks to the gneisses. 

The grain size of this basic suite varies greatly. Some textures arc those 
of the very coarse noritcs, others are doleritic, hut the average is that of a medium- 
grained norite. The composiiion varies somewhat from a charuockihe basic 
dionte to a typical norite. In hand-specimen Lhe diorites are similar to those of 
Mount WoodrofTe, but a little darker. They are also comparable microscopically. 
In hand-specimen the noritcs. however, arc coarse dull brownish-black heavy 
rocks composed mainly of large dull grey plagioclases, poikihtically including 
numerous small greasy grains of pyroxene. A Ihitc-t winning is not obvious m 
hand-specimen. There arc several large clots of dull greasy dark brown pyroxene 

crystals. f „ , , 

Microscopically, these norites consist of over 60% basic audesme with the 
rest of the rock composed of hypcrsthene and diallage. whose properties are 
simitar to those of the Mount Woodroffe rocks. Iron ores are not plentiful. No 
fluorescing zircons occur in these rocks. 

W The atrial photographs (see footnote, p. 19S) surest that there is very Jifcd* 
KiicfsMc structure iu (Hose ror.U. 'Some of that suggested in the photographs f the 
north-west flank 61 Mount Woodrotfe may be linked with the profound mylonituution. 
known lo occur a^ng* fjta northern JbnK of thr* Mount. 


(2) The Upsan Dowxs Gkanjtic Rocks, 

Little work has been done in the Upsan Downs area. These medium-grained 
granites are characterised by large flesh-coloured feldspar crystals. Considerable 
crush has taken place, and in some localities beautiful crusb-augen gneisses are 
developed (e.g., the Pass one mile to the north-east of Victor's Well, and at 
Erlywanyawanya Rock Hole to the south-west of Mount Woodroffe). These 
rocks contain no fluorescing zircon. 

At the rock hole in Brock Pass a granite occurs among contorted and parlly 
''assimilated" gneisses, This granite shows fluorescing zircon, thus, although shown 
on the map by the symbol used for the Upsan Downs Granite, it may well repre- 
sent another epoch. 

(3) The Nokjte to the SonTu-WEfir of Tjatjajtja. 

These norites comprise a considerable belt on the- spur two or three miles 
south-west of Tjatjanja. The rocks from this locality vary in grain sisc, ranging 
from coarse norite to rnicronorite. Normally the rockn are mottled greasy, dark 
blue-grey medium-grained, and the abundance of a resplendent bronzy biotite is 
the most striking feature. [Noted by Jack (1915, p. 18) ]. 

The plagioclase shows prominent albite-twin striae. Under the microscope, 
half of the rock appears to consisr of an acid labradonte. It is often associated 
With the hypersthene poikilitieally. The plagiodase is an antipcrthiie. 
Hypersthene and diallage are present in equal amount. Magnetite is scarce, as 
also is some interstitial orthoclase microperthite. No fluorescing zircon was 

A small orthite-bearing pegmatite similar to that at Koli Koli (see p. ZDfej 
cuts these norites near the northern-most junction with the charnoekitic granite 
of the Tjatjanja spur. Thus alone suggests that these basic rocks: arc older than 
those of the Ernabella massif, 

(4) Tun Erxabella Granodtoritic Massif, 

This rock type occurs as a great meridional belt with Enmbella near the 
centre. Though the. belt is granitic around Tjatjanja to the south, the massif, 
taken as a whole, is probably granodioritic. 

The. map shows ''cupolas" of similar material in several other places, notablv 
among the gneisses :— 

(a) South and north-east of Koli Koli rock hofe to the west. 

(b) South of the Siouy Jump Up to the cast. 

(c) Bald Hill and Spinifex Hill further to the cast. 

It is regretted that owing mainly to difficulty of terrain and climatic condi- 
tions, many of the junctions between the. gneisses and the non-gneissic acid rocks 
could not always be studied as well as one would desire- They have been located 
on the map, however, to facilitate future investigation. 

The best junctions were found on the western side of the massif in the Mount 
Carruthers area near Emaheila, and at Alalka to the north. In these localities 
large sheets of fresh rock are exposed in the water-courses, and the study of 
xenoliths and other phenomena is facilitated thereby. The eastern junctions are 
not readily studied. Many of these occur beneath sandy valleys. The junctions 
in the Tjatjanja area are relatively broad zones, but xenoliths are not common. 

In the Mount Carntthers area the trend of the xenoliths in the Plrnabella 
massif is between N^^E, and N. 35" E. with an approximate dip of the rude 
xenolithic lenses of 70° to the east. Small aplitic veins are often found with 


identical dip and strike of the xenoliths, but the pegmatite veins are almost always 
trending W. 10° N. and dipping south 75 u -80°. The strike of the gneisses is 
usually north-south, hence these xenoliths are usually af an angle of approxi- 
mately 25° 10 the gneisses. 'JW* mav indicate some prc-crystalhsation movement 
of the non-gneissic 'igneous mass" relative to the gneisses, from which it was 
probably derived by granitisation. 

In die Alalia Kock Hole area> \cnoli:hs arc common. The trend is between 
North and N, 18° E., the usual being about N. 15° \l. No evidence of aplitic or 
pegmatittc activity was. found. 

hi ah localities where xenoliths could be studied, a great diversity of xrno- 
lithic rock types was apparent. The xenoliths which are most obvious are am phi - 
bolitic, button closer investigation partly assimilated elongated fragments of 
gneisses of all types outlined above are found to be the more abundant, Xe.nolirhs 
in the area south- we*t of Mount CarrUthers were traced to their probable equiva- 
lents /« situ in some instances. A common size for the lens-shaped xenoliths is 
about 20 cms. x 5 cmfti x 10 cms. Many larger ant! smaller ones were found. 

A feature of many of the acidic inclusions is the presence of biotitc which 
has developed throughout the tens Basic inclusions usually show a marked 
reaction rim of hornblende. It is hoped later to publish some microscopic 
observations on the xenoliths. 

The conspicuous paucity of pegmatites, aplttes and quartz veins indicate* the. 
special '"dry" conditions under which these post-gneiss hypcrsthene-bearing acidic 
rocks crystallised. 

The Rocks of the Ernahdla Massif, whether granitic or granodioritic arc 
very similar in most of their characters. In hand-specimen they are medium- 
grained, greasy bluish dark grey granitic, rocks composed of bluish-grey feldspars 
of medium size, small hluc-greyish glassy quartz irregularly scattered, and small 
ragged areas of greasy dark brown feTromagnesrans. In several localities large 
phenocrysts of plagiociase are present (up lo n cms. x 2 cms.), ;md these with the 
rest of the plagiociase usually show splendid albitc-twin striae. Phenocrysts of 
the pyroxenes are found at times. 

Under the microscope these rocks are holocrystallinc and more or less 
allotriomorphic granular. For the most part the ferromagnesians arc of somewhat 
smaller dimensions than the salie constituents (average of rock 1*5 mm. to 
2 mm. long). 

The plagiociase (50%) is a mid-aridcsine and well twinned. It is sometimes 
a little antipe.rthi tic. Orthoclase mieroperthite (about 20%,) contains roundel 
quartz inclusions (millet-seed type) in considerable numher. These inclusions are 
uncommon in the plagiociase, 

Quartz (15%) is ragged and interstitial except wIhmi included in the polaf.a 
feldspar. Mvrmekite is plentiful. The tongued character of ihe quartz (yo 
common in 'ihe gneisses) is an uncommon feature of these non-gnetsxic 
chamockitic rocks. 

Preferred orientation nf the dark minerals is not present K> atiy marked 
degree, cither in the field or in microslide. 

ilvperslhene is not so notably pleochroic as in most gneisses and rocks of 
the Mount Woodroffe area. A weakly pleochroic or non -pleochroic pale grccmsb- 
grey monoclinic pyroxene (probably a species of di-opside) is plentiful. There 
is no apparent age difference in the two pyroxenes. Apatite and magnetite arc 
plentiful. Hornblende occurs, hut usually in small amount and has crystallised 
after the pyroxenes. Hornblende is plentiful, however, in the cupola phases, for 
instance, at Spimfex Hill and Hald Hill 

rViotitc occurs as small Hanking crystals on some of the patches of magnetite. 

Zircon is plentiful and H tfJ Hw fluorescent type. 

Analytical and petrography dtffci tlOV* been collected, but are being withheld 
fnr Ihc present, ft should be noted thai this rock type has all the characters of 
a "normal igneous rock." It is hoped to show later the paHngenette relation of 
this "igneous, type" to the gneisses. 

Ill . PEfi M AT1TES 

Pourly defined .small pegmatites outcrop throughout the Ernabella plain 
among the charnockitic granndiorices. Only occasionally are. they found among 
the gms&k rocks. The Musgrave Ranges, as a whole, arc very poor in peg- 
matites, An intensive search was made for these, and specimens and notes have 
been fctken of each nn^ found. Ap'utes are much rarer. Most pegmatites ate 
usually only about, one foot wide, and less than three or lour chains long 1 . Their 
strike i£ almost always West 10° North* with a dip South 75°-SO\ \n places 
doicrite dykes are found cutting across these pegmatites. 

The minerals noted are set out in approximate order of abundance. Haematite 
is often in well crystallised masses, Biofitc is usually much contorted, and ragged 
books occur up to 7 cms. in diameter, T-arge zircons are often found embedded. 
No mnscovite has been found anywhere in the ranges. 

Hornblende — This mineral is the dominant mineral of many pegmatites, and 
has been noted as large black shining crystals up to 8 cms. in length. (*artic! 
occurs in some of the pegmatites as dull greasy reddish-brown rounded subhedial 
crystals up Lo 3 cms. across. 

Orthite (Allav.ite)— Occasional small pockets of large tabular crystals were 
noted in a few of the pegmatites of the Ernabella plain. The mineral is Jet black. 
and has no cleavage but a pitch-like concboidal fracture. Specific gravity is about 
,V5. and hardness roughly Cr5. Though too weak to affect the electroscope., a very 
Weak radio-activity was revealed after an exposure of up to three weeks on photo- 
graphic plates. Throug-hoitt the ranges only a few pounds of the mineral were 
obtained, though a thorough search was made. Analytical work h in progress 
ro use tins mineral in age-determination. Zircon is present in certain pegmatites, 
as black or dark brown euhedral crystals, up lo 2 cms. long. Though very weakly 
radioactive (about the same as the orthite), they do not fluoresce under tjltra- 
Vjotet light. This is rernarkahle, for the zircons of the parent rock, the 
charnockitic granodiorites of Ernabella fluorescence in orange. Preliminary 
investigations with autoradiography indicate that the fluorescence decreases 
regularly as the radioactivity and depth of dark brown increase. Apatite occur? as 
tiny ctihedral pale blue crystals, but is rare. Tourmaline was fouud in only a 
few cases, and is of the common black variety 

Among the gneisses true pegmatites are rarely found. At Koli Kolf Rock 
lfo!e atl interesting- but quite small hornblende pegmatite occurs, in which -nail- 
like crystals of orthite and an onmge-rluuresemg euhedral resinous if zircon" occur. 
Similar pegmatites at Irawanja and Arkalanja show neither orthite nor zircon. 

Work is in progress on the pegmatites. 

IV. Dor.j-:iiiTE Dykes 

Cutting all of the other rock types, but probably not far removed m age from 
the Ernabella massif, is a large suite of do.'erites, A common trend is a few- 
degrees south of west; with a dip to the south of fiO°-70 D , Another set is present 
near Ernabella, with an approximate- north-south strike and dip to the east nf 
25 n -40°. Similar dyke-rocks occur all the way to Umbyarra, about 140 miles to 
the east, and according to Dr. Jack, far to the south-east "into the Everard Ranges, 
and beyond. Dr. Jack observed (Jack, 1915. pp. 15 ; 18)iu these regions to the 
south-east, thai (he basic dykes cut the gneissic and non-gneissic rocks, but not 


those of the Adelaide Series (Proterozoic — the "Cambrian" of the earlier geolo- 
gists). These dyke rocks, as seen in the Mnsgrave.s, consist essentially of labra- 
dorite lathes, with hypersthene, diallagc -and olivine combining- to give the typical 
ophitic texture of dolcrite. Specimens from about one hundred dykes' have 
been collected for description. 


Most rock types of the Musgrave Ranges have been mylouiiized in places. 
A large collection of material showing transition types has been made for future 
work. Pseudo-tachylyte:; are well developed. 

Both gneissic and non-gneissic types may show a crushing and a rolling out. 
They are often found to grade through a crush zone of mylonitized augen- 
gneisscs to dense hallefhnta types. The gneisses so formed are not to be confused 
with the gneisses outlined above. The crushed rocks are only in association with 
major fault zones. 

Though considerable areas of crushed rocks may be of later origin, most of 
this mylotiitissatiou seems to have taken place after the pegmatite? were emplaeed, 
and is probably associated with the widespread cpidotization noted with the joint- 
ing which led to the dole rite intrusions. 

The coincidence of the west-south-west trend of the dolerite dykes and major 
zones of mylonttization is evident in the following cases. 

Intense mylonitization (with pseudotachylytcs), cpidotizalion and basic dyke 
injection arc plentiful throughout: — 

(1) the tectonic valley stretching from Uusan Downs through the valley 
just north of Tjatjanja and Mount Ferdinand to the Gilpin's Well and 
liig Hill localities; 

(2) the tectonic val?ey from Taljaritja Well through Top Springs and along 
the creek south of One Tree Hill; and 

(3) the great tectonic valley running along TietkhVs Creek, through Nalja- 
Avarn, Ernahella, Rrock Pass and into the Oowallinna area. 

There is also a coincidence of these same features with the weaker develop- 
ment of the north-south basic dykes in the Ernabella. area. 

Tt is apparent that the conditions which allowed typical mylonitcs to form 
must have been considerably different from those prevailing- at the depths where 
the charnockites crystallised. Hypersthene is always absent from these mylonittzed 
rocks, and hornblende and garnet with bi'nrite and marked mortar structure are 

The Musgrave Ranges consist of a complex series of many rock types, all 
of Precambrian age. Present indications are that the chronological order of' the 
major rock groups seems to be the following: — 

1. Gneisses showing sedimentary origin and represented by garnet, cordierite, 
spinel and sillimanite gneisses, and containing no fluorescing zircon. This 
group is thought to represent the oldest rocks in this area. 

2. llypcrsihene-bearing Gneisses not showing obvious sedimentary origin but 
containing no fluorescing zircon. 

3. Wyperstlmtc-bearing Gneisses xot showing obvious sedimentary origin 
but containing a fluorescing zircon. 


4. Hypersthene-hearing non-tfncissic Intermediate and Basic Rocks of Mount 
Woodruff c and Trudinger Fuss. .These have no fluorescing zircon. 

5. The Upsan Datvns Granific Rocks. These have no fluorescing zircon. 

6. Hypcrsthcjie-bcuring non-gneissic Granites and Granodiarites of the Ernahcila 

These "igneous" rocks contain fluorescing zircon and xenoliths of the 

7. Pegmatites of the Ernabclla area. 

8. Mylonitizafion and Basic dyke injection. 

The hypersthene-bcaring rocks may be called chamockites. The position in 
the above sequence of the numerous altered basic dykes and masses of several 
epochs has yet to be determined. 

The gneisses of group 2 are thought to represent, in part, the pranilized 
equivalents of the ancient sediments. The gneisses of group 3. with their fluoresc- 
ing zircon, are tentatively considered to represent possibly a superimposed meta- 
morphism (with the introduction of considerable new material) of some of the 
earlier gneisses. This is well seen at Kuna-unpunja, where patches of tht rela- 
tively coarsely-grained and more granitic rocks of group 3 cut across acid, inter- 
mediate and basic types of group 2. Similar directional features are always present 
in both groups, but those of group 3 are almost always much less marked. 

A criterion suggested for the differentiation of the gneisses of group 2 and 
group 3, therefore, rests on the critical problem of the origin and introduction of 
the fluorescing zircon. The non-gneissic igneous rocks of group 6 may indicate 
a relatively mobile and somewhat contaminated equivalent of the gneisses^ of 
group 3, because these, as stated, contain plentiful fluorescing zircon and xenoiiths 
of all types of gneiss. The rocks of group 4 and 5 contain no fluorescent zircon 
and are thought to have crystallised after those of group 3. 

The ubiquity of hypersthene, the relative rarity of hornblende and biotite and 
the marked paucity of pegmatites signify the "dry" conditions under which these 
charnockitic rocks finally crystallised. 


I wish to express my indebtedness primarily to Professor Sir Douglas 
Mawsou, and also to Messrs. A. \V\ Kleeman and H. E. E. Brock of the Geology 
Department of the University of Adelaide, for interest and advice in many 
matters. D. R. Bowes kindly made two chemical analyses. Without the willing 
co-operation of the Superintendent (the late Rew J. Love), stafT, and natives of 
the Presbvterian Mission Station at Ernabella, the field work during the two 
seasons would have been impossible. Mr. Trudinger, in particular, rendered 
valuable hospitality, Finally, I must thank R. Brock for his outstanding assist- 
ance while accompanying me on the second expedition to the area. 


Basedow, II. 1905 ''Geological Repent on the Country Traversed by the South 

Australian Government North- West Prospecting Expedition, 1903." 

Trans. Roy. Soc. S. Aust M 29 
Basedow, H. 1915 " Journal of the Government North-West Expedition 

(March to November 1903). Proc. Roy. Geog. Soc. Australasia, 

S. Aust. Branch, 15 

Vans. Roy. Soc. S. Aust., 1947 

Vol. 71, Plate II 

Fig'. 1. Looking north-north-west from Naljawara, 3' miles east of lumahella. 
Note how island-like hills of non-gneissic charnockitie granodiorite punctuate 

tine plain. 

* ■?• 

Fig. 2. North-north-east approach to Trudinger Pass, taken two miles south-wesi 

of Kuua-unpunja. Most of the rocks are of gncissic charnockite. with norite in 

the j>ass. Tjakunja on left; Landmark Hill on right in distance. 

Fig, 3. Mount Spec, taken from northern junction of mica norite to south-west 
of Tjatjanja. Most rocks are acidic and intermediate gneissie charnockiUs. 

Fig. 4. Non-gneissic charnockitie granodiorite cut by parallel doleriU- dykes, 
South-west tip of hills, two miles west of Palpatjara Well. 

Trans. Roy, Soc. S. Aust., 1**47 

1 \ * » t«i 

Fig. 1. Looking nortli from Mount Carruthers, showing north-south gnelssic 
charnockitcs in distance on left, Top Springs and Ernabella Creek gorge in centre 
in distance, and Mount I tjinpiri and other non-gneissic eliarnoekitic granodioritc 

area's to the right 

Fig. 2. Looking north from east side of Razor Hill, nine miles north-north-east 

of Krnabella. Razor Hill consists of non-gnefcsic charnockitic granodioritc; hills 

to right are gtteissic and in part show obvious sedimentary origin. 

Trans. Roy. Sec. S. Auet., 1947 

Vol ?), ['tnti l\ 

i'\u. I. ).n->kinM rasl iViiin cumtnit .if Miftiiit Wogdroffc Cover 5.000 Feet, hfjibtst 

mountain in South Australia). Mount Spec i- flat-tapped motintaiti mi Un in 

distance A stunted specter pi "fipirufcx' 1 (Tritfdiai is ubutHlant'. 

i ti . 1. Looking wc.-t ir^m ?inun)il of XT mint Wpad rafifo. Foreground • 

'ii-. .,!■.,, 1 ilir .|ii:m i .- liu.rilf \\itll £iu'i — ir granitic d ..m,,,, lit- 

\l a [fir vi*:-;rt.-ili^n -tiiniol ;pi .'i. * | ij "I ri(;i_ 

,i u ii 
i fatrbt. 

a, wasox 

Trans. Roy. &QC. S. Aust, 71, (2), Dec. 1947 

26* 15' S, 

OMBAGUNOA **b* .*. * 

( # blackmill 

i r. 



26' 3C 




GEOLOGY • Central *E 



1' , • '.| GNEISSIC CHARNOCKITE (oldest rocks) 




r — SJ BASIC DYKES (post charnockite) L_ 




/ta^ ItoU**^ **«k 

mm m 


Brown, H. Y. L. 1890 "Report on journey from Warrina to M-isgrave Ranges." 
Part. Pap. No. 45 "* 

Carrutiikks, J. 1892 "Triangulation of North-west Portion of South Australia." 
Pari. Pap. 179 

"Elder Exploration Expedition, Scientific Results of" 1893 Trans. Roy. 
Soc. of S. Aust., 16 

Forrest, j, 1875 "Explorations in Australia," London 

Giles, E. 1872-1874 "Geographical Travels in Central Australia, 18724874" 

Gosse, \V. C. 1874 "Reports and Diarv of Mt W. C. Gosse's Expedition in 
1873." Pari. Pap, No. 48 

Hubbe, S. G. 1897 "Stock Route Expedition from South to West Australia." 
Pari. Pap, No. 51 

Jack, R. L. 1915 ''The Geology and Prospects of the Region to the South of 
the Musgrave Ranges, and the Geology of the Western Portion of the 
Great Australian Artesian Basin." Geological Survey of S. Aust " 
Bull. 5 

Maurice, R. % 1904 "Extracts from Journals." Pari. Pap. No. 43 

Murray, W. R. 1901 "Report and Map." Pari. Pap. No. 148 

Wells, l* A., and George, F. R. 1904 "Reports on Prospecting Operations in 
the Musgrave, Mann and Tomkinson Ranges." Pari. Pap. No. 54 



By Reg. C. Sprigg 


A richly fossiliferous horizon has been discovered within the massive Pound Quartzite formation 
which underlies the Cambrian Archaeocyathinae limestones in the Flinders Ranges, South 
Australia. The fossils occur as impressions on surfaces of flaggy quartzite. The five genera and 
species described are almost certainly all pelagic Coelenterates, and while several forms are referred 
to the class Scyphozoa, it is possible that one or more species may be more correctly assigned as 
Hydromedusae. The more problematical forms may prove to be pneumatophores or swimming 




By Reg. C. Sprigg * 

[Read 8 May 1947] 




A richly fossiliferows horizon has been discovered within the massive Pound 
Quartzite formation which underlies the Cambrian Archaeocyathinae limestones 
in the Flinders Ranges, South Australia, The fossils occur as impressions on 
surfaces of flaggy quartzite. The five genera and species described are almost 
certainly all pelagic Coclen derates, and while several forms are referred to the 
class Scypbozoa, it is possible that one or more species may be more correctly 
assigned as Hvdromcdusae. The more problematical forms may prove io be 
pneumatophores ov swimming bells. 

In this paper a group of fossils recently discovered in the uppermost forma- 
tion of the Adelaide Series (Upper Proterozoic to Lower Cambrian) is discussed. 
The fossils were found on a rise 
approximately three hundred yards 
south-west of the principal south- 
mine workings at Ediacara, near 
p.eltana. South Australia. They 
occurred as impressions in flaggy 
quar^i;^ and are among the oldest 
drect records of animal life in the 

i I the environmental associations 
of the forms have been correctly 
interpreted there is good reason to 
consider all the forms pelagic and free 
swimming. They all appear to lack 
bard parts and to represent animals of 
very varied affinities. All are pro- 
bably Coclenterates and all may be 
jeHytishes, although in at least two 
cases insufficient detail is available to 
make reliable comparisons with any 
living or fossil animals (vis., Papilio- 
vcUi, Dickinsonla and the unnamed 
circular form described in association 

with Beltanella). Just possibly the latter two forms may be floats or pneumato- 
phores of colonial coelenterates. 

One form (Ediacarja) is referred tentatively to either of the orders 
Semaeostomeae (Discomedusae) or Rhizostomeae of the class Scyphozoa, while 
another {Beltanella) may be referable to either of the classes Hydromedusae or 

♦Assistant Government Geologist, South Australia. 

Trans. Roy. £bft S. Aust., 71, (2), 1 December l!>47 



Fossil jellyfish as casts- and impressions have, been described from the Lower 
Cambrian of New York, Sweden, Russia r.nd hlobemia, and now from Australia; 
from the mid-Cambrian of Britisli Columbia and Alabama; from tbe Silurian of 
Victoria and from tbe Permian of Saxony; from the Jurassic of Solnhofen. 
Kavaria and from the Cretaceous of Texas. The identity of tonic of those 
fossils has been questioned, and some, are better referred to die Problematica. The 
stratigraphic range of jellyfishcs is almost certainly pre-l'ambrian to present. 

As indicated above the fossil jelly fishes described in this paper are very 
probably Lower Cambrian in agft The fossil impression* -were found in the 
"I'ound' qtiartzitc formation of tht Upper Adelaide Series. This massive forma- 
tion, which frequently measures several thousands of feet in thickness, imme- 
diately precedes massive Cambrian Pleospongia (— Archaeocyathinac) lime- 
stones. The impressions occurred at a horizon approximately 100 feet strati- 
graphically below the base of the Hinttlone and approximately 600 feet below the 
first Pleospongia remains yet located at this locality/ 1 * 

Generally speaking the horizon of the Pleospongia is considered to be- 
near the top nf the Lower Cambiian (David, 1927). On this assumption the 
present fossils would have been living in about middle Cower Cambrian rime. 
The "Pound" quarlzite in which ihe fosftih were found [j considered temaiivclv 
(IVbnvson, 1939) to represent the base of Cambrian sedimentation in South Aus- 
tralia, although there is reason to believe that the dawn of the Cambrian may 
eventually be taken back still further (Sprint*, VM2), 

With perhaps the exception of David's (1936) Ispahan problematic^, and 
certain annelid tracks common in several quartsti'tcs of the Adelaide Series, the 
new fossils represent the oldest undoubted marine animals recorded in Australia 
(tig-. 2). They provide the first reliable indication that the Pound puaru. to is 
of marine origin. 

Considering the extremely perishable nature of jellyrtsbes and related soft- 
bodied coelenterales (many jellyfish contain 99 fc sea water), ft is remarkable that 
any of them should have left traces of their existence in the fossil state. It is 
obvious that very special conditions pf burial would be necessary for their pre- 
servation. Walcoit (1898) suggests thai in tbe case of many United States 
Cambrian forms the medusae probably had the habit of living- on a muddy bottom 
hi great numbers. Associated fossils suggest a shallow water environment. There 
appeared to be rapid burial and consolidation of the sediment, not by exposure 
between tides, but entirely beneath the water. Tn other cases, such as at Soln- 
hofen. Bavaria (Lower Jurassic), medusae have been found in extremely fine- 
grained slates. 

Agassi* {1862) notes that the living- Aurrh'a ftavida, afler the spawning 
period, is frequently seen in large numbers floating on the water. There has been 
a thickening of the tissues by an increased deposition of animal substance, The 
disc of the animal has become thin and almost leathery and it is more clastic (and 
at the same time more brittle) than before. Many of the marginal appendages 

(l > It is to be noted t!iat, contrary to popular conception, the South Australian 
Pleognongia are not reef builders. Their ?patial distribution within the enclosing 
sediment simulates "coral meadow V conditions. This explains why. in this 
ca^-e (and in many other cases) Archaeocyathtnae do not occur right from the 
base of the so-called Archaeocyathinae limestone. The Arcnaeos are usually restricted 
tn more or 1*^ definite horizons within ihn limestone formation, and even in the T&GSi 
highly tossiliferous of these, actual foc-vil remains form only a relatively small portion 
o£ the rock. They are not massed remains as in modern coral reefs. 


of ihe umbrella and oral region drop off during this period. It can be seen that 
in this dried out condition the medusae, when stranded on beaches and covered 
by sediment, oiler much better chance for fossilizatiom 

Caster (1945) notes that when Aurelia and other medusae are washed ashore 
and stranded during low tide in midsummer, they quick'y dry out on the surface. 
"Dehydration of the aqueous jelly brings out in surface relief embedded struc- 
tures, which in life would hardly be discernible, except by transparency, on the 

exterior. While the upper surface 



S70 * 


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I I 

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sapor* s&s&srs. 

fb£s/?£t 5as?^$to/?e 


Generalised Section across Cambrian and 
associated formations of Flinders Ranges 
[ modified after Sir D. Mawson 1939 (2) ], 

showing approximate horizons of known 

is hardened, thus inhibiting de- 
hydration, the surface in contact 
with the heuch remains soft and 
often turgid as in life. Depending 
on the texture of the strand, the 
imprint made by the surface may 
he faithful or indistinct. The in- 
coming tide picks up the partially 
embedded jellies and carries them 
further ashore, and occasionally 
turns them over to embed them 
aga : n at turn of tide with the erst- 
while downside up." He notes 
further that extraordinarily rapid 
solidification of the entombing 
matrix is not necessary, as Evrii 
jellyfish can be dug from tidal sands 
of today wherein they have appar- 
ently remained buried for a much 
longer tune than tidal periodicity. 
In the ca?e of the newly dis- 
covered fossils we arc dealing 
exclusively with buried pelagic 
forms. The fossils axe impressions 
in flaggy sandstone quartzite, and 
the grooves of the impressions are 
stained with lerruginous material 
or possess* a film of clayey material. 
The enclosing sediment was origin- 
ally a fme-grahied and well-sorted 
sand which had accumulated near 
the western margin of the vast 
Flinders sedimentary geusynciiue. 
The environment of entombment 
was that of intertidal Hats or o£ the 

Toughening of the surface tissues 
of the fossil animals as described by 
Agassiz and Caster seems almost 
certainly to have occurred prior to 
burial, and judging from the atti- 
tude of the fossils in the field 
the animals were mostly preserved 
with their ex-umbrella surfaces 


It is noted that the fossils were all found on "free" faces of flaggy quartzite. 
and always on the upper surfaces of these slabs. (Beds dip at 10° to 20°). The 
fissility of the quartzite appears to have been controlled to some extent by the 
presence of clayey films, which might also have had significance in the preserva- 
tion of the fossil impressions. 





Genus Ediacaria Sprigg, gen. now 

Genotype Ediacaria flindersi Sprigg, gen. et sp. nov. 

Pound Quartzite, Upper Adelaide Series (Lower Cambrian), 
Ediacara, South Australia. 

Being monotypic the new genus shows the species characters described below. 
Generic characters include the hell-like manubrial structure and other structures 
nf the central disc area. Comparisons between Ediacaria and other most closely 
related genera are given below, 

Ediacaria flindersi Sprigg, gen. ct sp. no v. 
(PI. I, fie- 1) 
llolotype: No. T.l, Tate Museum Collection, Adelaide University, South Aus- 

Description — Medusa impression circular, radially symmetrical ; surface 
flattened, but with radial and concentric features of low relief. Three concentric 
zones are clearly distinguishable. 

Inner Zone — (?) Manubrium bell-like, constricted near its junction centrally 
with the sub-umbrella surFace and expanded distally. It lies over sideways and 
is compressed laterally. Length 15 mm., and maximum width (flattened) 14 mm. 
At least three pendant lobate pouches extend 9 to li mm. centritugally from the 
base of the manubrium. Beyond these pouches the central zone is essentially 
smooth, although there is an incomplete concentric groove half-way to the zone 

Median Zone — Surface smooth, somewhat inflated : zone delimited on inner 
and outer aspects by concentric grooves — one (or two) on inuer margin, and one 
deeper with associated minor and less regular grooves on the outer. Two well- 
marked radial grooves are present, while indistinct radial striations are more 

External Zone — Surface flattened or only slightly convex in transverse 
section with minor concentric undulations or flirtings and numerous radial grooves 
or striae. In the annular segment reprcsenling three-fourths of the perimeter, at 
least -14 separate radial grooves can be recognised. Although somewhat irregular 
in themselves, they are distributed around the zone relatively evenly. Most diverge 
centritugally, but some converge in this direction. The outer margin (perimeter 
of fossil) is fairly regular (circular), and with one or two doubtfnl exceptions 
is devoid of marginal notches. A concentric groove lies approximately 4 mm. in 
from the perimeter of the form. 

< 2 > Classification used in this work is based on that of Parker ant! Haswcll 1940. 


Dimensions — Largest diameter 114 mm. Respective widths of inner, median 
and outer zones along greatest radius 20 mm., 17 mm. and 25 mm. 

Discussions and Comparisons — The specimen is considered to be the impres- 
sion of the sub-umbrella surface of a ''dried out" jellyfish. Organs adjacent to the 
oral surface of the original animal have come lo stand out in relief, and the 
manubrium stands out strongly. The central /one probably corresponds with the 
gastrovascular cavity, and external structures of the central disc region are .super- 
imposed upon it. 

The sub-triangular ( ?) manubrial structure has been so interpreted because 
of its apparent fusion centrally with the. sub-umbrella surface, and because no 
other comparable structures are distributed radially about the centre. The flattened 
attitude of this manubrial bell bears a superficial resemblance to the insert lobes 
of the central d:scs of Kirkhrndla (Caster) and Rhizostow-ifes (Haekel). How- 
ever, the absence of more of these structures radially disposed about the centre 
tersely contradicts this view. In lite the manubrial structure would be suspended 
verticallv from the central region. The shape of the mouth opening cannot be 

&?&'/&/ C^tiv'*: 

+uw»-\ •'-"-'•'-'A 1 /-', pft I '. Xn, '.'. I ' ! ! ! ! TFrnmsp **%,/££* 

Fig. 3 

Ediacwla fl.nilcrsj. 

A, details oi the subumhrejto Mirfaec of the reconstructed form; B r cross section 
through radial canals; C, actual profile section along a diameter of the fossil. 

The three pendant pouches extending - radially from the base pi the manubrium 
are possibly gastro-genhai sacs in connection with radial canals. Judging from 
the distribution of tile three sacs preserved there were probably eight of these in 
the original animal. 

Various concentric (lutings, with the exception of that adjacent tbe margin 
of the form, are referable to "the circular muscles of the sub-umbrella. The epi- 
margmai groove is probably a circular canal, in which case the narrow flange 
beyond it would probably be a "hood. 1 ' 

The well-marked radial grooves of the median zone correspond with inter-, 
ad- s or per-radiai canals, whereas the much finer, numerous radial striations of 
the outer zone and to a much lesser extent of the median zone suggest more minor 
canals, splitting* and radiating towards the circular canal. This may merely be 
shrinkage creasing, hut in any cases such creases would tend to follow stich rela- 
tive weaknesses as the. canal lines. The grooves are Rub-para1iel and tend to 
increase in number centrifugally, a feature which is in keeping with the canal 


Two marginal notches can be interpreted ; these are at intervals corresponding 
with the separation indicated by projection to the margin of the stronger radial 
canals of the median zone. It is noticeable that in each case deeper radial 
Mriations continue to each notch The annular (?) hood structure, where observed 
beyond one marginal notch, is indented m sympathy. This would support the 
view that the notches arc regular marginal features, possibly originally enclosing 
sensory structures. On the other hand it is noted that in other portions of the 
fossil where continuous sections of the margin are preserved, other notches are 
not apparent. This would suggest that the two notches noted may be accidental 
invaginations of the margin consequent upon deformation during burial. 

There are no indications of marginal tentacles, hut in view of the 
nature of the enclosing sediment and the probable delicate nature of such tentacles 
tf present, it is difficult to imagine that evidence of them could have been pre- 
served. Furthermore, experience with modern jellyfish (Aurtiia, etc.) indicates 
that in many cases the marginal tentacles drop off in the senescent, stages. 

There appears little doubt that Ediacarki is a Scvphozoan. The form 
obviously had a flattened saucer or disc-like umbrella, and for this reason is 
referred to either of the orders Semaeostomeae (Discomedusae) or Rhizo- 
Ntomcae. To decide further to which of these orders the form belongs, a detailed 
knowledge of the structure of the mouth and oral arms would be necessary. In 
view of tjje nahire of the preservation of the specimen this cannot be hoped for. 
However, it is noticeable that the manubrial structure as interpreted is relatively 
simple — a fact which suggests correlation with the Semaeostomcac* or could per- 
ceivably indicate an even simpler class relationship (vis., IJydrozoan), The 
absence of marginal tentacles on the. other hand is a Rhkustomid character, hut 
in view of the difficulties already pointed out evidence such as this is primarily 
negative. It docs seem, however, that no strong tentacles existed — a fact which 
would preclude membership of the order Trachymcdusae of the class Trachylinae. 

Detailed comparisons with other fossils is exceedingly difficult in view of the 
lark of knowledge of many critical features, but it is noticeable that closest (super- 
ficial) resemblance is perhaps with F<hbostomiics and Semaeostomites (both 
Haekel) of the Upper Jurassic of Solnhofen, Bavaria. In these forms three 
concentric zones can be inferred, but otherwise there is little similarity in avail- 
able detail of the central disc regions. Ring muscles arc well developed in the 
outer portions of Rhtsostomites^ as they are in Ediaearia. No obvious ring canal 
is present in Rhbostomites as it is in Ediaearia and Semaeosiomiies, and whereas 
the margin of Scmaeostomites is split up into 120-128 marginal lobes, such sub- 
division is not apparent in the other two forms. 

Ediacaria(?) sp, Sprigg 
(Pi. i, %. 2) 

Specimen No. T.2.. Tate Mn\. Coll., Add. Univ.,. S. AusL 

Description — Impression fragmentary; original form apparently circular, but 
less than one-eighth of the perimeter is preserved, although the outline of the 
central disc region is almost complete. In all, three concentric zones can be 
made out. 

Central Zone — Surface smooth except where badly preserved; slightly 

inflated ; perimeter sub-circular. 

Median and Outer Zones — Zones poorly differentiated ; median otic is smooth 
and of variable width. Outer zone inflated somewhat towards outer margin, 
;tnd traversed by numerous sub-parallel radial grooves which fade rewards either 
margin. Exterior margin slightly lobate with indications of two minor notches. 


Discussion and Comparison — In view of the absence of structures in surface 
relief in the central and median zones of this fossil, it is suggested that here is a 
cast of the ab-nral or ex-umbrella surface of a jellyfish. The poor state of pre- 
servation of the centremost portion of the fossil form prevents complete assess- 
ment of this fact, but nevertheless there appear to be no signs of mouth or gastro* 
genital structures. The centrally inflated region appears to have collapsed over 
the stomachs of the animal and preserved, therefore, the roughly circular outline 
of this region. 

In the outer zone which, presumably, in the living animal was thin, the sub- 
parallel radial grooves are well preserved. These are very numerous and appear 
unbranched; they may represent radial canals or merely he shrinkage creases. 
Prominent radial striations continue directly to the two minor marginal notches* 
and there are no signs of marginal tentacles. 

Tentatively the fossil is considered to be an impression of the ab-oral surface 
of a species of Ediacaria. As with the holotype specimen, three concentric zones 
can be interpreted, the outermost of which is traversed by very numerous radial 
striations. Indefinite notches which are separated by approximately equal intervals 
appear marginally in hoth forms. Width relations of the respective zones oi the 
fossils agree favourably. 


Genus Bcltanella Spriggj gen. nov. 

Genotype Beltanella gilesi Sprigg, gen. et sp. nov. 

Pound quartzite, Upper Adelaide Series (Lower Cambrian) Ediacara. 

South Australia, 

Being raonotypic this new genus shares the species traits described below. 
Until more is known of the fossil, generic characters should include the octagonal 
arrangement of the circular (?) gonadial structures and their association with the 
radial canals, the presence of a well-developed delicate peripheral umbral or velar 
structure and the simple circular oral aperture. 

Beltanella gilesi Sprigg, gen. et. sp. nov. 
(PI. % fig, 1) 

Ilolotypc: No. T.3., Tate Mus. Coll.,, Adel. Univ., S. Aust. 

Description — Medusa impression circular. Umbrella rather flat, but falling 1 
away sharply near its outer margin. (?) Velarium horizontal, depressed approxi- 
mately 4 nun. in relation to the fiat ex-umbrella surface. Umbrella region sub- 
divided into two zones by a faint annular groove as follows. 

tvner Zone — Surface smooth, broken only by annular grooves respectively 
3 and 12 mm. in diameter at the centre. Centremost area depressed very slightly. 

Outer Zone — Surface dominantly flat, but slopes away steeply near outer 
margin of umbrella. This secondary (sloping) surface has the form of a highly 
truncate cone whose apical angle is approximately 80 degrees. Zone characterised 
by the presence oi circular (?) gonadial structures, approximately 10 mm. in 
diameter. These regular structures are arranged on cither sides of the major 
radial canals in an octagonal pattern centrally within the zone. At least 
four of these Can be recognised and each possesses an inner concentric 
groove 3-4 mm. in diameter. Two paired radial grooves (? canals) are diametri- 
cally opposed, and a third set lies radially at right angles. The grooves pass inter- 
mediate between ( ?) paired gonadial structures but do not continue into the inner 



The ex-umbrella surface is slightly lobate at the edge of the flat raised 
portion, but below where the conical surface meets the (?) velarium the margin 
is smooth. 


■£*f-^mf STlr.^tr./ 


Fig. 4 

Beltanella gilesi 

A, details of exumbrclla surface and related embedded organs; B, transverse 
section through restored form; C t true profile section along diameter of form- 

Velarium — Structure marginal, obviously thin, well developed; unclulose 
surface depressed; undulations annular in plan. 

Dimensions — Maximum diameter of fossil 110 mm., minimum 97: widths 
along: single radii of inner and outer zones and velarium respectively 18-20, 21-23 
and 10-14 mm. 

Discussion and Comparisons — The specimen is the cast of the cx-umbrella 
surface (ab-oral) of a jellyfish. 

The central zone evidently corresponds with the gastrovascular cavity as at 
its margin it gives off paired grooves which are interpreted as interradial canals. 
There are no signs of subdivision within the cavity and no indication of com- 
plicated manubrial structures. The simple circular grooves situated centrally may 
be oral structures, or possibly representative of a collapsed truncate gastric 
cone which occurs in some jellyfish to aid in the even distribution of food to 
various portions of the animal's stomach. 

The radial grooves of the outer zone are thought to be interradial canals, 
although why they should be paired is not known. There is no sign of branch 
canals from them, uor is there present any groove suggestive of a circular canal. 
The circular (?)gonadial structures which are distributed evenly around the 
centre of this zone may be considered as paired in relation to the supposed radial 
canals. The central annular grooves of each gonadial structure may mark a genital 

The (?) velarium or peripheral umbral jelly is remarkably well preserved 
considering its obvious delicateness ; its contained annular undulations may indicate 
ring muscles. 

The decision to place Belt&neUa within the Scyphozoa must be regarded only 
as very tentative, as the animal has many restrictive features characteristic of 
either certain orders of the Scyphozoa or of the Hydrozoa. For example, the 
simple mouth, the presence of a few unhranched radial canals and the association 


of the gonads with the radial canals are Trachylinid (Hydrozoan) characters. On 
the other hand the flattened disc-shaped umbrella, its relatively large size, and the 
absence of large tentacles arc more characteristic of the Scyphozoa. It appears 
very likely, therefore, that Beltanella is a member of a group, ancestral to cither 
or both the Scyphozoa and Hydrozoa of modern times. Indeed, this is to be 
expected considering the great age of the fossil. The author knows of no living 
or fossil jellyfish with which useful comparison of the foregoing specimen can 
be made. 

The fossil is tentatively classified with the Scyphozoa, mainly in view of the 
absence of the large tentacles which are typical of the Trachylinids. 

On the slab of rock carrying the holotype there are at least four additional 
circular structures (10-23 mm. in diameter) which may represent juvenile forms. 
All display annular grooves, and in the largest specimen, the central portion (9 mm. 
in diameter) is raised relatively to the outer zone. In the latter example (which 
is photographed with the holotype) there is very definite evidence of a velar 
structure, 2 mm. wide, similar to that of Bclianell& 

Still another possibly related form found at the same locality is the disc-like 
impression shown on pi, ii.. fig. 2 (specimen No, T4). This external cast is almost 

Fig. 5 
True proiile section of un-named form figured on pi, ii, fig. 2. 

perfectly circular with a slightly raised rim near the externa! margin. The whole 
structure stands in relief a little above the face of the quartzite slab, and its 
surface is evenly convex, except in the marginal region, where it is raised some- 
what; it shows no recognisable anatomical structure or ornamentation. This 
smooth depressed (annular) area (7-10 mm. wide) is faintly reminiscent of a 
velar structure. Maximum diameter is 104 mm., not including the latter structure. 
The impression may be that of a simple discoid jellyfish or of a hydroid float. 

Class (?) HYDROZOA or (?) SCYPHOZOA 

Genus Cyclomedusa Sprigg. gen. now 

Genotype Cyclomedusa davidi Sprigg, gen. et sp. nov. 

Pound Quartzite, Upper Adelaide Series (Lower Cambrian) Ediacara, 

South Australia. 

This genus also is monotypic, and in view of the lack of detail of most of 
the critical features of the form the generic characters are based tentatively on 
the sculpturing of the ( ?) ex-umbrella surface of the form. 

Cyclomedusa davidi Sprigs gen. et sp. nov. 
(H. i% fig. 1) 

Holotype; No. T.5., Tate Mus. Coll., Adel. Univ., S. Anst, 

Description — Impression circular, depressed, with concentric undulations. 
Central portion raised, distinctly nodular. The whole form exhibits striking 
radial symmetry. Surface subdivided by at least seven annular grooves. For 
convenience three major zones are recognised. 

Inner Zone — Consists simply of the hemispherical nodular portion mentioned 
previously. Approximately 2*5 mm. in radial width and 1'5 mm. in height. 

Median Zone — Subdivided into two annular portions of low relief, the inner 
of which is traversed in turn by two annular grooves, while the outer is 


ornamented by very numerous radial striations about 3 mm. in length. There arc 
approximately 16 of these grooves in each quarter. A poorly developed annular 
groove traverses the outer portion. 

Outer Zone— Surface sculpture very similar to that of the median zone; 
the smooth inner portion is bounded by annular grooves and the outer is traversed 
by numerous poorly developed radial striations averaging 4 mm. in length. The 
outer margin to the zone is very poorly preserved. 

Dimensions— Maximum diameter probably 50_mm. Average widths of the 
inner, median and outer zones along single radius 2*5, 11*0 and 11-0 mm. 

Discussions and Comparisons— The surface ornamentations which stand in 
strong relief, appear to be of superficial significance, but some of the annular 
Outings may 'be related to ring muscles of the subumbrella of a jellyfish. No 
structures which can be attributed to a body cavity or to gonads are visible. 

Cyclnmi'dusa davidi 

A, details of the surface ornamentation of the 
restored form; 

B, profile section along a diameter of the 

There is no evidence of extra-marginal structures, but as noted, the marginal 
region of the fossil is poorly preserved. 

In view of the paucitv of critical detail it is only with hesitation that the fossil 
is provisionally included within the Coelcnccrata. The form is highly problematical 
and possibly only represents the restricted central portion of a larger animal. It 
does seem certain from its regularity and complexity, however, that the form is 
not a pseudo fossil. 

Genus Dickinsonia Sprtgg, gen. nov. 

Genotype Dickinsonia costata Sprigs gen. et sp. nov. 

Pound Quartzite, Upper Adelaide Series (Lower Cambriau) Ediucara, 

South Australia 

This genus which is monotypic, exhibits the species characters described 
below. Animal of ovoid form, '(f) inflated aborally and possesses a marginal 
crenulate flange. Median longitudinal furrow gives off very numerous subradial 
grooves to the outer (crenulate) margin of the form. 

Dickinsonia costata gen. et sp. nov. 
(PI. iii, fig. 2) 

Holotype: No. TA, Tate Mm. Coll., Adel. Univ., South Australia. 

Description— Impression ovoid, bilaterally symmetrical, essentially flat. 
Median longitudinal furrow approximately 35 mm. long, gives off 80 to 90 radiat- 
ing- or diverging grooves or costae (?) alternatively to the outer margin of the 
fossil. Margin slightly crenulate, the notches corresponding with the intersection 
of the radiating grooves. Well developed concentric epi-marginal sulcus marks 
off a marginal flange. 

Dimensions— Length 68 nun.; width 60 mm. Flange width variable, from 
2 to 7 aim., due to distortion. 


Discussion and Comparisom—Tht fossil is the impression of the (?) dorsal 
aspect of a bilaterally symmetrical animal of very doubtful affinities. During 
burial the animal was flattened and compressed slightly obliquely in a manner 
which suggests that it was strongly convex dorsallv. The animal Was symmetrical 
across both longitudinal and transverse planes. Radiating grooves or costae may 
represent chrtinous rods or canals, while the epimarginal groove mav represent 
a circular canal No gastrogenital structures or appendages are apparent. 

It is exceedingly difficult to classify Dickinsonia on the little detail available. 
The author knows of no related animal with which to establish relationships, and 
until new specimens with more detail are found little more can be suggested* than 
that the animal is probably a coelenterate. 


Fig. 7 

Dickinsonia cvstala 

Details of surface ornamentation of the restored 

form as viewed in elevation (A) and in plan (B) 

There are certain superficial features in which it resembles the problematical 
form Discophyllum peltatum (HaJI), but these apparent resemblances soon dis- 
appear when detailed comparisons are made. Discophyllum is a rounded or oval 
flattened form with bars which radiate from the centre of the form, and not from 
a longitudinal furrow as in Dickinsonia, Also there are fine concentric undulating 
lines that cross both the rays and the interspaces between them. Comparable 
structures do nof occur in Dickinsonia. There is no flange structure in Disco- 
phyllum, nor is the external margin crenulate. The ribs fade out before the 
margin is reached. 

(?) Dickinsonia sp, 
(PI. iv, % 1) 

Specimen No. 1.7. , Tate Mus. Coll... Adel. Univ., South Australia, 

Description — Impression fragmentary, radially costate; costae continue to the 
margin. Margin rounded, slightly crenulate in sympathy with the disposition of 
the costae. A shght bending of the costae, noticeable 7-8 mm. in from the margin 
of the fossil, indicates a tendency to flange formation. 

Discussions and Comparisons— From the little detail available for comparison 
of this specimen with the foregoing holotype the major similarities are concerned 
with the radiating costae, which fit both forms are strongly developed and continue 
to the somewhat crenulate margin. In the holotype' specimen the costae arc- 
interrupted by a very well marked concentric sulcus, whereas in the latter specimen 
there is only the faintest hint of interruption of the radiaJ costae in a similar 
position. None of the costae in the second specimen appear to diverge as if to 
unite alternatively into a central furrow as in the type form. 

The correlation of this fossil with Dickinsonia is quite tentatve and the form 
must be considered problematical. 


Genus Papilionata Sprigg, gen. no v. 
Genotype Papilionata eyrei gen. et sp. nov. 

Pound Quartette, Upper Adelaide Series (Lower Cambrian) Ediacara, 

South Australia. 

The genus is mnnotypic and shares the species characters described below. 
Unfortunately, the generic characters must be based on the shape of the fossil 
and its restricted surface sculpture. The form is bilaterally symmetrical, 
papilionaceous with the amygdaloidal "wings" fused anteriorly. A marginal 
groove is present and the posterior rnargiu of the wings is met by sub-parallel or 
slightly radiating grooves. 

Papilionata eyrei Sprigg, gen. et sp. nov. 
(Pi. iv, fig. 2) 
Holotype: No. T.8., Tate Mus. Coll., Adel. Univ., South Australia. 

Description — Impression bilaterally symmetrical. Left hand portion im- 
perfectly preserved and fragmcntal. Complete impression papilionaccaus, the 
•'wings'' being fused for approximately 30 mm, in the anterior aspect. The left 
hand wing is amygdaloidal in plau, curved convexly on the inner margin, and 
rounded and slightly lobate (posteriorly) on the lateral margin. From the 
posterior portion of this margin, radial grooves converge towards a central point 
near the inner margin, but fade out after 10 to 20 mm. The grooves are stronger 
and longer at the posterior extremity, and they are not visible forward of the 
centre of this margin. The single notch on the lateral margin may not have 
anatomical significance, being probably a crenulation due to the animal's assumed 
position of rest. The lateral margin is paralleled by a well-developed groove 
through which the radial costae continue uninterrupted. The groove diverges 
slightly from the margin at the anterior end. A similar divergence is apparent 
in the "reflected" wing. 

Dimensions — Maximum width of complete extended form 1 50 mm. 
Maximum length and width of single wing 112 and 56 mm. respectively. 

Discussions and Comparisons — The impression has been described as bi- 
laterally symmetrical but there is a possibility that two separate organisms may 
be represented, the line nf "fusion" as described being a fortuitous overlap of the 
respective organisms. Nevertheless, the excellent ''reflection" of the two portions 
appears to predispose of this view. The single wing impression suggests 
a disc-like jellyfish lying upon its margin with portion of its umbrella surface 
folded under. 

The author knows of no similar organism with which useful comparison 
can be made, 


The author is indented to Dr. Curt Teichert for valuable suggestions at 
several stages in the preparation of the manuscript, and to Professor J. A. Prescott 
for undertaking the photographing of the fossils at the Waite Research Institute. 

Acassjz, L. 1862 "Contributions to the Natural History of the United States 
of America," 4, 63 

Castek, K. E. 1945 "A New Jellyfish (Kirklandia texana, Caster) from the 
Lower Cretaceous of Texas." Palaeontographica Americana, 3, No. 18 


Chapman, F. 1926 "New or little known Fossils in the National Museum." 
Pt. xxx : A Silurian Jellyfish. Proc. Roy. Soc. Vict., 39, (1), 13-17 

David, T. W. E. 1927 "Note on the Geological Horizon of the Archaeo- 
cyathinae." Trans. Roy. Soc. S. Aust., 51 

David, T. W. E, and Tillyard, R. J. 1936 "Memoir on Fossils of the late 
Pre-Cambrian (Newer Proterozoic) from the Adelaide Series in South 
Australia." Angus and Robertson, Sydney 

Ktesunger, A. 1939 "Scyphozoa" (in Schindewolf, O., Handbuch der Palaeo- 
zoologie), Bd. 2 A, Lief 5, Berlin, 70-109 

Mawson, D. 1939 (1) "The Late Proterozoic Sediments of South Australia." 
A.N.Z.A.A.S., 24 

Mawson, D. 1939 (2) "The Cambrian Sequence in the Wirrealpa Basin." 
Trans. Roy. Soc. S. Aust., 63, (2) 

Parker, T. J., and Haswell, W. A. 1940 "Text-Book of Zoology." Mac- 
Millan & Co., London 

Sprigg, R. C. 1942 "The Geologv of the Eden-Moana Fault Block." Trans, 
Roy. Soc. S. Aust, 66, (2), 201 

Walcott, C. D. 1898 "Fossil Medusae," Monograph. U.S.G.S., 30, 

runs. Roy. Sue. S. AiM.. 1 ( M7 

VuK7l, I'lak V 

Fig. 1 Ihiiaicaria flindcrsi Sprigg 
(apiirox. x-S). 

Photo1»y EC P. Phillips 

l ; i^. Z lldiacarm (?) flindo'si Spngg 

SliulitU less llum natural size. 

Photo l)j K I". Philh> 

Trans. Rov, Soc. S. Aust, 1947 

Vol.71, Plate VI 

ig, 1 Bcltam'tki jii'csi Spring 
Two thirds natural size. 

Pllotnhj K. P; PlttlHpa 

J' III, 2 1 >i-i-til<l ScVlJho^>ail iU" /noit); 

J'raiiN, Kt»\. Soc S. \ust« 1'M! 

Vol.71, i'lau VII 

Im#. I Cyvhntetfyjtti t/cffic/i Sprigs i .-. I3) 

l'lin 1 hj K. I 1 , I'liil'ii'- 

!-ii-]iiK in -< 1 than natural -.i/r. 

I- Ity K 1". I'hilliiw 

Trans, kov. Sue S. Au>i., 1947 

Vol.71. Plate VI If 

Fig, I Pit kiiisoiiitt (?) s/-. (apprax, xlii 

Pliotafcy K. P. Phillip* 



i ig, 2 Pafifwuata cyrci Spffgg- 
i approx. n .; ). 

Phutnby K. P. Phillips 



By J. W.Evans 


A collection of Homoptera recently sent me by Mr. D. C. Swan included representatives of eight 
species of Jassoids belonging to the family Eurymelidae, which he had collected in New Guinea 
during the war. Five of these species are described below. Of the remainder, one, Eurymeloides 
nigra Evans, is confined to New Guinea, and two, Eurymeloides punctata Sign, and E. adspersa 
Sign., both of which are well-known Australian species, are recorded from New Guinea for the first 
time. The host plants were species of Eucalyptus and Melaleuca. 



By J. W. Evans* 

Communicated by D. C. Swan 
[Read 7 August 1947] 

A collection of Ilomoptera recently sent mc by Mr- D. C. Swan included 
representatives of eight species of Jassoids belonging to the family Eurymclidae, 
which he had collected in New Guinea during the war. Five of these species are 
described below. Of the remainder, one, Eurymeloides nigra Evans, is confined 
to New Guinea, and two, Eitrymeloides punctata Sign, and E, adspersa Sign., both 
of which are well-known Australian species > are recorded from New Guinea for 
the first time. The host plants were species of Eucalyptus and Melaleuca, 

Of the remaining four new species at Eurymelids descrihed below, the most 
distinctive is Ipoella daznsi, which was collected in North-west Australia by the 
late Dr. Consett Davis, whose tragic death in 1944 was an irreparable loss to 
entomology in Australia. 

Eurymeloides sogerensis sp. nov. 

Length, 5 mm. Head, face evenly dark brown, but for the outer margins of 
the lora and maxillary plates, which are cream. Pronotum brown mottled with 
pale greyish-brown. Scutelltwi dark brown. Tegmen dark brown, irregularly 
mottled with dark grey. Thorax, ventral surface and legs, dark brown. Male 
Genitalia, aedeagus as in fig. 1, A. 

Holotype, J, from Sogeri, New Guinea (D. C. Swart, 5/45), in the British 
Museum. Paratypes in the South Australian Museum. 

Eurymeloides motuana sp. nov. 
Length, 7 mm. Head dark nigger or reddish-brown mottled with yellow; 
antc-clypcus dark brown; lora and adjacent parts of the maxillary plates 
yellowish or pale brown. Pronotum nigger or reddish-brown mottled with ivory. 
Scutellmn, the angles black, centrally dark brown with oval yellowish markings. 
Tcgmen black with two irregular white, transverse fasciae and with a white 
marking at the distal apex of each anal vein. Legs brown, tile bases of the 
spines and the proximal tarsal segment of the hind tibia, white, Male Genitalia, 
aedeagus as in fig. l r B< 

Halotype, S , from Port Moresby, New Guinea (D. C. Swan, 5/45), in the 
British Museum. Paratypes in the South Australian Museum, 

Eurymeloides nigrobrunnea sp. nov. 
Length, 7-2 mm. Head, maxillary plates pale brownish-yellow { ante-clypeus 
lora and fronto-clypeus pale reddish-brown; vertex dark brown mottled" with 
apricot yellow. Pronation and ScuUilum chestnut brown or brown mottled with 
yellow, Tegmen black with two white fasciae, each with a large irregular black 
marking, anal veins distally white. Legs, femora pale brown, tibiae dark brown, 
the bases of the spines and the proximal tarsal segment, white. Male Genitalia, 
aedeagus as in fig. 1, C. 

Holotype, S , from Port George, North-west Australia (C, Davis, 5/43), in 
the British Museum. Paratypes in the South Australian Museum. 

* Imperial Institute of Entomology 

Trans. Roy. Sac. S. Auat„ 71, (2), 1 December 1947 

Ipoides brunomaculata sp. nov. 
Length, 5 mm. Head, face, but for the vertex, yellowish-ivory with a median 
longitudinal dark brown stripe. Vertex pale brown irregularly mottled with 
chestnut brown, Crown and pronation pale chestnut brown mottled with 
yellowish-ivory. Scutellum deep chestnut brown with two pale semi-circular 
markings. Tegmen pale hyaline brown with irregular white markings; veins 
white, brown in part. Legs very pale brown. Male Genitalia, aedeagus as in 

fig. i.'f. 

Hohtype f 8 , from Port Moresby, New Guinea (D. C. Swan, 5/45), in the 
British Museum, Paratypes in the South Australian Museum. 

Ipoides melaleucae sp. nov. 

Length, 5 mm. Head, face, maxillary plates greyish, the rest of the face pale 
greyish-brown evenly mottled with dark brown. Crown, pronotum and scutellum, 
greyish, sparsely mottled with brown. Tegmen very pale hyaline brown; veins 
brown with white bars. Legs, femora dark brown, tibiae pale brown. Male 
Genitalia, aedeagus as in fig. 1, G. 

Holotype, 3, from Sogeri, New Guinea (D. C. Swan, 5/45), in the British 
Museum. Paratypes in the South Australian Museum. 

Anipo pallescens sp. nov. 
Length, 5*5 mm. Head, face, maxillary plates, lora and fronto-clypeus 
laterally,* ivory partially suffused with apricot; ante-clypeus, the fronto-clypeus 

medially and the vertex, deep coffee brown with pale oval markings. Crown and 
pronotum pale greyish-brown mottled laterally with dark brown. Scutellum, 
anterior lateral angles dark brown, the remainder pale brown. Tegmen pale 
hyaline brown with evenly distributed circular colourless hyaline areas. Thorax, 
ventral surface and legs marked with a pattern of pale and dark brown. Male 
Genitalia, aedeagus as in fig, 1 , E. 

Halolype, $, from Sogeri, New Guinea ( D. C. Swan, 5/45), in the British 
Museum. Paratypes in the South Australian Museum. 

Ipoella davisi sp. nov. 
Length, 5 mm. Head, face almost flat ; ante-clypeus narrowly depressed 
anteriorly, dark brown; fronto-clypeus medially dark brown, shading to pale 
brown, laterally ivory; lora brown adjacent to the ante-clypeus. laterally ivory; 
maxillary plates ivory anterior to the antennae, dark brown beneath the eyes; 
vertex chestnut and very dark brown, posteriorly ivory. Crown brown with 
irregular dark brown markings. Pronotum huff mottled with brown. Scutellum 
dark brown. Tegmen hyaline, the cosial margin and the apex smoky brown, the 
remainder dark brown with a broad proximal white fascia and white* markings at 
the apex of the first anal vein. Some specimens have also a distal white fascia. 
Male Genitalia, aedeagus as in fig. 1, D. 

Holotype, 3 , from Jsdell River, Walcott Inlet, North-west Australia 
(C Davis, 9/43 on Ficus), in the British Museum. Paratypes in the South Aus- 
tralian Museum. 

Bakeriola tasmaniensis sp. nov. 

Length, 5 mm. Head, face black sparsely mottled with brown but for the 
outer margins of the maxillary plates, which are white and the lora, which are 
cream, partially suffused with pale brown. Crown of head and pronotum pale 
brown mottled with dark brown. Scutellum dark brown. Tegmen, clavus con- 
colorous with the pronotum, the remainder hyaline brown with circular and 
irregularly shaped colourless area. Legs dark brown but for the coxae, the tarsi, 
and both ends of the tibiae and femora, which arc pale brown. Male Genitalia, 
aedeagus as in fig. 1, H. 

Holotype, $ from Risdon, Tasmania (J. W. E., 4/39), in the British 
Museum. Paratypes in the South Australian Museum, 

Bakeriola rubra sp. nov. 
Length, 4*8 mm. General coloration dark brown. Head, face evenly convex, 
dark brown mottled with reddish-brown. Crown and pronotum irregularly 
mottled with dark brown and reddish-brown. Scutellum dark brown, mottled 
antero-mcdially with pale reddish-brown . Tegmen, hyaline dark and reddish- 
brown with oval white markings. Legs reddish-brown, hind tibia whh three spurs 
and several spines. 

Holotype, <£ , from Moolooka, Queensland (4/44, E. F. ftiek), in the British 
Museum. Paratype i n the South Australian Museum, 


Trans. Roy. Soc. S. Aust., 71, (2), Dec. 1947 








J Banded Siliceous Limestones 

H Slates & Shales (Tapley Hill Series) 

G Tillite Series 

x Arkose Grit in Tillite Series 

F Sub-Glacial Quartzite 

E Slates Phyllites & Hornfds 

---j C Slates aPhyllites with somShales& thin beds of Dolomite Limestone 
B Quartzite & Sandstone 
A Phyllites & Slates with some thin beds of Dolomitic Limestone 

v Alluvium with no rock outcrops 



By J. W.Evans 


Kangaroo Island lies off the South Australian coast at the base of Gulf St. Vincent, being separated 
from Fleurieu Peninsula by Backstairs Passage (10 miles wide) and from Yorke Peninsula by 
Investigator Strait (about 26 miles wide). The Island is 90 miles long and up to 32 miles wide, 
narrowing to only Vi mile wide between the American River inlet and the south coast (see fig. 1). 
The long axis of the island is approximately east-west; the island lies between latitude 35° 5' S. and 
35°34 1 / 2 ' E. and 138° 8' E. 



JBy H. B. S. VVoMI.KFiEY* 

(Rwid 7 August 1S»47] 

Kangaroo Island lies off the South Australian coast at the base oi <iv\i 
St Vincent being separated from Fleurieu Peninsula by Backstairs lassage 
(10 miles wide) and from Yorkc Peninsula by Investigator Strait (about 2o miles 
wide 1 ) The Island is 90 miles long and up to 32 miles wide, narrowing to only 
A mile wide between the American River inlet and the south coast (see tig. I). 
fee Ion? axis of the island is approximately east-west; the : island .lies between 
latitude W 5' S. and 35° 344' S, and between longitude 13b 32 E. ,mrt 
1 ^S 3 R' V 

The situation of Kangaroo Island in relation to the mainland, and the shape 
of the island itself, result in great variation in conditions of roughness tftfm the 
const The exposed and rough south and west coasts contrast markedly with the 
calmer areas of the north coast, while the American River tidal inlet forms a 
distinct tvpe of habitat not found elsewhere around the island 

From the point of view of algal ecology Kangaroo Island offers a particularly 
satisfactory area for study, especially in illustrating the control exerted on the 
algal flora by the degree of wave action. 

This paper is the first report on work carried out during the past tour 
vear*. The aim lias been, firstly, to give a general account of the mterlidul algal 
ccotoey of a part ot the Southern Australian coast, since there have been no 
previous ecological studies of this region; secondly, to obtain as comprehensive a 
list as possible of the species present; and thirdly, to carry out autecological 
studies of the more characteristic and dominant species. 

In this paper it is proposed to describe the more important environmental 
factors for the island as a whole; to discuss the terminology found most satis- 
factory and to Rive a preliminary general account ot the broader aspects ot the 
aleal ecology. In subsequent papers the more detailed ecology of characteristic 
regtoris will be dealt with, and a census of the known species will be given. 
Floristic and ecological comparisons with other areas will also be left to httei 
papers By first presenting a general survey of the algal ecology of the whole 
island, it is hoped to give perspective to the later detailed descriptions ot indi- 
vidual localities. a a 
The localities around the island which have been studied are shown m ng. 4. 
Of these, Pennington Bay and American River have received most attention- It 
has been possible to pay only one or two short visits to the western end of the 
Hand while the very rough nature of the country and lack of roads prevent--, 
visits to most of the intervening parts of the south and west coasts. This, and 
the restriction of field work to the University vacations, have limited examination 
Of most localities other than American River and Pennington Bay to January. 
Seasonal changes in places other than American River and Tenmngton Bay are 
therefore unknown as vet. but the associations which occur in these places and 
ore described in this paper are almost certainly present throughout the year. 

* Department of Botany, University of Adelaide. 

Tr^tn. R«y- Src. S. Aust., 71, t2>. I December 1947 

The areas studied at Western River and Middle River consist of the coast 
on cither side of the river mouth. These rivers run only after heavy rain, at 
other times being blocked by a sand bar at the mouth. American River, however, 
is an extensive tidal inlet, consisting of a series of lagoons with Avide tidal flats 
and a central channel, opening to the sea through a mouth some. 250 yards wide. 
The amount of fresh water entering the lagoons from small creeks is negligible, 
except for possible local efTects after heavy rain. The conditions at all "River" 
localities, therefore, are truly marine. 

Previous Marine Ecological Stwjes in Australia 
Australian phycology h based to a large extent on the work of the early 
European algologists, Harvey, Agardh, Sonder and others, who described collec- 
tions sent to them from Australia. As a result, very little ecological information 
ie available about the early localities from which collections were made. Harvey 
(1854), however, gave short notes on the Western Australian coast, and ecological 
information about many species in his Phycologia AustraUcn* 

More recently, Hedley (1915) has presented a very general survey of the 
New South Wales coast, and Lucas (1935) has described the algal ecology of 
Lord Howe Island. Pope's (1943) survey of animal and algal life on a reef near 
Sydney is the only work of a detailed nature, but deals primarily with the animal 

The review of Chapman (1946) gives a comprehensive list of references to 
algal ecology studies elsewhere in the world. 

Previous Records from Kangaroo Island 
Previous records of marine algae from Kangaroo Island arc very few 
CIdand and Rlack (1941) listed 11 species collected from near the mouth of 
Sou 1 West River, and determined by A. H. S, Lucas, Lucas (1929), in his 
census, lists two species from the island, while Tart 1 of "The Seaweeds of South 
Australia*' records Caalerpa hedlcyi W. v. fiossc, "dredged in some 8 fathoms off 
the coast." Part II of the "Seaweeds of South Australia" contains several 
records., all incorporated by the present author. Two short reports on the 
Rmdanas and a new species of Dasyopsis have also been published (WomeHey 
1046 a and b). 

*r T Mo la ^ d ve " etatio11 of Ka ngaroo Island has received considerable attention. 
Tate (188o) gave the first general account of the Island, and more recently 
Wood (1930) has elucidated the relationships of the flora and shown a high 
degree of endemism, particularly in species confined to the western end, Baldwin 
and Crocker (1941) have described vegetation communities in the central part uf 
ihe Island. "^ 


1. The Coastal Geology 

The present study has shown that while the individual type of rock has 

little effect on the flora, the rock topography, in that it may result in different 

types of habitats, may be of considerable importance in determining the algal 

associations of an area. 

The only reasonably detailed geological map of Kangaroo Island is that of 
Wacc (1915), which is followed in fig. 1. The backbone of the island consists 
of Pre-Cambnan schists and gneisses, overlain in the central part by siliceous 
sand and latente. On the west coast, eastern part of the north coast, atid to a 
ksser extent on the east coast, these rocks form magnificent cliiT scenerv The 
coast west of Emu Bay is of later age ( Post-Cambrian ) (Madigan 1928 K 



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West B. 


The most significant geological features of the south coast of Kangaroo 
Island are the outcrops of ancient rocks at the capes. These outcrops do not 
rise to any notable height ahove sea level, and mostly appear at the foot of cliffs 
which fringe, the shore. The extent of these outcrops of Prc-Cambrian rocks »3 
shown in fig. L Two types of rocks, granite arid quartzlte or mica schists, form 
alternating patches along some of the coast. Between the areas of older rocks 
the coast consists either of sandy beaches backed by sand dunes or of sand- 
rock cliffs and horizontal reefs formed from older consolidated sand dunes. This 
rock weathers into very sharp edges and pinnacles, and by its variable hardness 
makes an irregular coast. 

The two different types of rock found on the south coast of Kangaroo Island, 
iVv the flat rock platforms of consolidated sand-xnek such as at -Pennington Bay, 
and the harder more steeply sloping rocks .such as seen at Cape Willoughby, west 
of Vivonne Bay, and Cape Coudie, result in two very different habitats for algal 
and animal growth, and appear to illustrate the conditions occurring along most 
of the south coast (cf., pi. ix, fig. 3 r and pi. xii, fig. 1). 

Most of the American River inlet consists of extensive sandy or sandy-mud 
tidal flats, but in many areas, particularly in Pelican Lagoon, these stretch out 
from low cliffs of the same consolidated sand-rock as at Pennington Bay on the 
south coast. 

The northern coast of the island is composed mainly of ancient rocks sloping 
off into 3 to 12 or more feet of water, giving a relatively small area of rock- 
exposed at low tide. 

2. Decree of Wave Action 

The degree of roughness of any locality is of prime importance in determining 
the algal associations present. The south and west coasts are exposed without 
any protection to the Southern Ocean, and conditions are invariably rough. Heavy 
breakers are a constant feature of this coastline (see pi. ix, fig. 1). Passing 
along the north coast, from west to east, conditions become progressively calmer, 
owing to the shelter afforded by the mainland. Northwards from Cape 
Willoughby and past Hog Bay wave action is moderate, with breakers only in 
tough weather. American River and Tetican Lagoon, quite unlike the rest of 
the island, form nn almost land-locked area where wave action is at a minimum. 

Although wave action is of great importance as an ecological factor, its 
measurement in any satisfactory way seems impossible. Average values of the 
forces from wave action to which algae are subjected in any one locality arc 
needed. These should be measured over short periods (when extremes may 
uccur) J as well as over monthly and yearly periods. In the absence of ajiy such 
measurements it is necessary, m some cases, to use the algae themselves as an 
index of the conditions. Thi* has hern done in subdividing the Rocky Shore 
Formation into subtormations, depending on the presence or absence nf 
Cystophora intermedia J. Ag. This alga is dominant in the sublittoral fringe on 
rough rocky coasts, but is replaced by other species nf Cystophora on calmer 

3. Tides 

The tides around the Australian coast, including the main characteristics 
and ranges of the tides in the South Australian Gulf region, have been described 
briefly by R t W. Chapman (1938). The tides around Kangaroo Island are of 
the semi-diurnal type, with two maxima, one appreciably lower than the other, 
and two minima during each 244-hour period. Fig. 2 "shows the form of the 
spring and neap ("dodge") tides at American River. 


1800 2200 

200 600 

TIME (Hours) 



Fig. 2 

Tide curves for spring and dodge tides in the American River Inlet and at 
Pennington Bay (spring tides only). The range of each tide curve and times of 
high and low water are comparable, but the heights given for each are arbitrary. 
The curves are derived from 24-hour surveys carried out at the American River 
Jetty, Muston Jetty, (3 miles south of American River Jetty), Pig Island in 
Pelican Lagoon, (2 miles East of Muston) and at Pennington Bay on the 
following dates: spring tides Jan. 10-11, 1947; dodge tides Jan. 16-17, 1947. 


South Australian tides vary so greatly from piace to place along the coast, 
both in their nature and times of high and low water, that it is necessary to 
obtain actual records from each locality. The tides have been analysed at com- 
paratively few places, especially around Kangaroo Island. An automatic tide 
f*auge recently established at Hog Bay should give most interesting results when 
records become available. The data given below for Kangaroo Island tides are 
derived from information made available by (he Soulh Australian Harbours 
Hoard and trom 24-hour surveys carried out at Amerxaa Kivcr and Fennioe;ton 
May. U will be evident that until accurate and more expensive tidal data are 
available, general limits and heights only can be given for the main algal acmes.. 

Tidal raTige around Kangaroo Islaurl is small. Along- flic BOU th and west 
CWste the spring range is about 2> feet. Passing eastwards along the north 
coast it increases to 4£ feel at Kingscwte, just over 4 feet at American River and 
Hog Bay, 4£ feet at Antechamber Bay, while a rise of h fee; is recoided from 
Cape. Willoughhy from old data (but this is probably too high). The wap (or 
"dodge") tides probably have a range of about 1| feet on the south roast and 
24 feet on the north. On the south and west coasts rfic smalt tidal range means 
that the wind and strength of the swell may exert nearly a% grcai an "effect as 
the tide ii^elf, and little reliance can be placed on the tidesalone" 

The most notable peculiarity of South Au*tra1 : an 'ides 5s rhe "dodging'* 
tide. This is discussed by R. W. Chapman (1924). At l J ort Adelaide, where 
the effect k most prominent, the water level may remain almost constant for 
24 hours or more at the neap periods. The cause *is that during the neap period 
the sun and the moon, together with the other tide-producing fences, exert almost 
equal hut opposite effects, one nullifying the other. It has been suggested that 
the abnormally large effect of the sun is accounted for by the synchronising of 
(he natural period of swing of the basin of water between Australia and 
Antarctica with the period uf the tide-producing forces. 

At American River (see tide curves, fig. 2) the dodge effect seems to be 
present, though small, at the neap period. For about 6 hours the water level 
remains almost stationary, before the next rise ot fall commences. Along much 
of the north coast this period of steady water level seems to occur, but no data 
are available as yet apart from isolated surveys at American River. 

During winter the mean sea level at Port Adelaide is. from 4 inches to 
h inches higher than in summer. This applies also tn Kangaroo Island, with con- 
sequently higher ndes during the winter mouths. With heavy west to north 
weather during winter very high tides often occur along the north coast. This 
is due to lnvesiigalnr Strait being about £j times as wide as Backstairs Passage, 
with consequent building up of the water mass in the area north of Kangaroo 
Island under the influence of westerly weather, The higher sea level during 
winter is of considerable importance on the south coast, where the increase is 
layge compared with the tidal range. The level of the horizontal rock platforms 
pi I he Pennington Bay region appears to correspond approximately with an 
average low neap tide level in summer. While north winds and low ltdes occasion- 
ally leave much of the reefs exposed m summer, whh consequent drying and 
desiccation of the algae, this rarely, if ever, occurs in winter. Apart from 
allowing a heavier growth in winter, this is also one of the factors controlling 
seasonal changes on the reefs. Similar consideraiions mav applv to a less extent 
in the American River inlet. 

4. Currents 

The surface current flows frnm west to cast across the Great Australian 
Higbt, passes along both sid*s. of Kangaroo Island and on towards Tasmania 


(see "Australia Pilot/ 1 1, 24). Eastward from the Bight the coastal current is 
strongest in the period May to July, with an average rate of 7 miles per day off 
the coast between Kangaroo Island and Cape Northumberland. During February 
to April and August to October it is weaker, averaging 3 miles per day, while 
from November to January it averages less than 2 miles per day. The currents 
in Backstairs Passage are largely tidal, reaching a speed of 2\ knots. 

5. Temperatures 
Observations taken around the coast of the island give the following results 
for sea temperature {Table 1). No data from other sources are available, but 
the figures agree well with the temperature isotherms given by Sverdrup et al 

Table I 

Sea Temperatures around the Coast of Kangaroo Island 

South coast— Summer (Jan.) - - insliore 19-20° C, offshore probably 18° C. 

Winter (early June) - inshore 16° C. 

(late July) - inshore 13*5° C. 

(Sept.) - - inshore 14° C. 

North coast— Summer (Jan.) - - inshore 20-21" C> offshore 19-20° C. 

Winter (June) - - inshore 11-13° C, depending on depth and 

air temperature; offshore 13-14" C, 

From this table it is evident that the yearly range of sea temperature on the 
south coast is small, being about 4° C, offshore and 5 to 6° C. on reefs. The 
range is greater on the north coast and depends greasy on the depth of water and 
degree of roughness, since the calmer water is affected much more by air tem- 
peratures. On the tidal Hats at American River temperatures as high as 32° C. 
have been recorded during summer in 6" - 12" of water, and as low as 10° C in 
winter. Algae on the flats must be able to withstand a far greater range in tem- 
perature than south coast forms. In isolated rock pools at Vivonne Ray and 
along the north coast temperatures of up to 30-34° C. are frequent in summer. 

Aiu Temperatures 
The climate of Kangaroo Island is fairly uniform. Some data for Kingscote 
arc given in Table II. The humidity figures probably give little indication of the 
humidity near algae exposed at low tide. Air temperatures are of greatest 
importance when a hot day (sometimes 35 - 3S° C. in summer) coincides with a 
low tide. Under such conditions algae on the Pennington Bay reefs may be 
almost or quite exposed for several hours and considerable damage may result. 

Table II 

Air Temperature and Humidity Data for Kingscote 
(From data made available by the South Australian Weather Bureau) 

All readings were taken at 50 feet above sea level, over a period of 17 years. 

Jan. Feb. Mat-. April May Junt Jt\\y Adr. Sept. Oct. Nov. De£. Av. 

Mean Max. 
Temp.°C. 22-3 22-7 21-4 19-3 17-7 14-9 14-2 14-5 15-S 17-8 19-8 21-4 18*4 

Mean Min. 
Temp.°C. 14-5 L5-6 14-2 12*5 11-0 9-6 8-7 8-5 9-3 10-S 12-1 13-8 11-7 

Mean Rcl. 
Humidity 70 74 75 76 80 83 82 .80 79 75 72 72 76 


6. Salinity 

Chlorinity of sea water on the south and north coasts is within the range 
19*6 - 19'9°/ 00 (salinity 35*4 - 35 -0%^). North coast values are usually slightly 
higher than those from the south coast. At Pelican Lagoon chlorinity in sum- 
mer reaches 20'5°/ O0 (salinity 37*0"/ on ), while in isolated rock pools (some with 
a heavy growth of Enteromorpha) summer chlorinity figures of 24*0%)o have been 
obtained. After three days of heavy rain (January 1946) chlorinity on the tidal 
flats at American River decreased to 17 Q f w ; such conditions, however, are very- 

Normal sea salinity around Kangaroo Island is high compared with other 
regions (<\gr., 34*9°/ o salinity near Sydney (Pope 1943) ). 

7. Phosphate anu Nitrate 
Estimations of these two major nutrients are as yet too few in number for 
any general conclusions to be reached. It appears, however, that nitrate is often 
extremely low (less than 1 part per 10°), while phosphate is rather variable. 
Phosphate figures of 14 and 23 p.p. 10 !t have been obtained from the south coast, 
and values between 2 and 60 pp. 10" from the American River inlet. Isolated high 
figures obtained at American River are prohahly due to the large bird population. 

8. Alkalinity 
The pH of water (by colorimetric methods) at Pennington Bay is about 
fr2^&*4i while at Pelican Lagoon figures of 8'1 have been obtained. 

9. Dissolved Oxygen 
The constantly boisterous seas on the south and west coasts result in water 
supersaturated with oxygen. The water on reefs at Pennington Bay is usually 
about 110% saturated with oxygen. Oxygen figures in shallow water at the 
American River inlet in summer show high supersaturation during the day 
(120- 150%, rarely a> high as 250% ) ? dropping to 50-70% saturation at night 
(an extreme of 10% saturation has been recorded). Such large ranges are due to 
the heavy growth of algae in the calm shallow water. For the most part it is 
unlikely that oxygen content of the. water is of importance in the algal ecology. 
The Winkler method was used in all estimations. 

10- Li:ght 
No attempt has been made to measure light intensities at different depths, 
but correlation of shaded littoral areas with communitics_ of sub-littoral algae lias 
been observed at Pennington Bay and Vivonne Bay. On the south coast, with 
constantly broken water, light penetration will be less than in calmer waters off 
the north coast At American River the large amount of silt carried in the tidal 
current reduces light penetration and may influence algal distribution. 


At the present stage there is little uniformity in nomenclature used in marine 
algal ecology. Chapman (1946) reviews opinions expressed about terminology, 
and advocates adoption of the terms used in land ecology. However, land ecolo- 
gists are far from agreement on their terminology, and until many more marine 
ecological studies in different parts of the world have been carried out, uniform 
and satisfactory meanings of the terms cannot be expected. This applies par- 
ticularly to the Australian coasts. As in the past, each worker must use the 
terminology which best suits his locality and his own concepts. 


The definitions adopted here have been found satisfactory in describing the 
algal vegetation of the Kangaroo Island coasts. Only further studies in other 
regions of the Southern Australian coast will show to what extent the concepts 
need to be modified, and their usage at present makes no pretension to be final. 

Association and Community* 

The concept of an association is fundamental to all ecological work, yet nidny 
different meanings have been applied to the term. The association is used here 
in the sense of a grouping of organisms distinct m species composition and fades 
from another grouping. It is composed of a dominant or dominants usually 
accompanied by other species whose presence is determined by responses to 
factors similar to those influencing the dominants (see Rees 1935)* This concept 
is to some extent subjective, but experience shows that most associations are 
objective entities. Studies over a long stretch of coast are usually necessary 
before the associations present can be determined. Intensive work on\small area's 
often results in variations of one basic association being considered as separate 
associations. On the other hand, an association may be scattered in its occurrence 
and cover areas of only a few square feei of rock, yet may be typical o\ that 
particular habitat, and pure and well defined in its occurrence. This is especially 
true of irregular and dissected coastlines. 

When associations occur during certain periods of the year only, ihcv arc 
classed as "seasonal associations. *' 

The term ''community" is commonly used with the same meaning as associa- 
tion, but often in a more general sense. It is applied in this and following papers 
when the status of the algal grouping has Hot been satisfactorily established. 


Apart from classifying the associations in their zones (see later), the onlv 
other grouping used is the formation. This is applied to the principal types of 
marine vegetation, much as it was used by Cotton (1912) for Clare Island and 
Rees. (1935) for Lough fne. Cotton's formations were based on the substrata 
and environment for algal growth, and comprised the following: 

1. Kocky shore Formation. 

2. Sand and Saudy-mttd Formation. 
3- Salt-marsh Formation, 

lie also distinguished: 

4. Vegetation of river mouths. 

5. Vegetation of brackish bays. 

Of these, the Rocky Coast j^ii&Ortf 1 of Cotton) and Sand and Sandv-mud 
hormations are found around Kangaroo Island, and they are real and natural 
entities. Rees' formations are based to a larger extent than Cotton's on the 
degree of wave action, but this is nearly always closely associated with the nature 
of the coast, and there is little difference between the formations of Rees and 
Cotton. The distribution of algae around Kangaroo Island shows that the degree 
of wave action is the most important environmental factor, as was emphasised 
by Rees. 

Use of the substrate and environment as criteria for "formation" is criticised 
by Chapman (1946, p. 658), who advocates following the practice of naming 
UMid plant formations on the dominant species. However, Tansley (1940'), to 
whom Chapman refers, describes Salt-marsh and Sand-dune Formations, anrf the 
naming of formations on the type of plant, while conveying at the same time some- 
thing of the nature of the environment, is common m land ecology (<>.</, mallee, 


savannah woodland, forest formations), Naming of the formation nn the 
dominant plant or animal species is quite impracticable in many cases, such as with 
the Sand and Sandy-mud Formation. 

In any case the formation is an abstraction. Of the four chief characteristics 
of an association, z>iz. } floristic composition, life-form, strucLure and habitat, 
Tansley uses two only (life-form and structure) as a basis for uniting associations 
into formations. It is just as logical to use eilher habitat or floristic composition 
as a criierion for such higher grouping (Ooxker and Wood 1947). 

Some algal formations to Tansley J s sense cati be readily determined by hfc 
form, e.g., a" blue-green formation and coralline-mat formation. On Kangaroo 
Island these formations are formed essentially of a single association, each 
delimited by definite environmental conditions, with the former occurring above 
the latter, 'it is evident that when the distinctive zones of algae around a coast- 
line have very different life-forms, one zone musl be chosen on which to base any 
classification; around Kangaroo Island this is the upper sub-littoral zone. To 
regard each zone as a distinct formation is clearly not justified, 

' It is well not to lose sitfht of the fundamental principle underlying the 
existence of communities, namely, that certain species live tog-ether in a particular 
situation because they have been selected by that environment, i>„ all of the 
species have the same habitat requirements for growth, This gives the ba*ts for 
defining associations. 

Within any one tidal zone, different associations may occur depending upon 
local variations*. These associations often have similar life forms, and possess 
units' in the fact that they have certain habitat requirements in common. Such 
"habitat zones" arc realities and are the natural units of higher grade than the 

In practice tide level, degree of exposure to wave action and nature 01 the 
substratum arc the chief habitat factor. Using he. latter two criteria a large 
unit (formation) is obtained; depending on the degree of wave action, the forma- 
tion may be divided into sub-formations; and by using tide levels zonmion is 
obtained. . .. 

Using habitat factors as criteria does present us with realities common m all 
parts of the world, and in this lies a real hope of achieving some decree oS 
uniformity in marine ecological nomenclature. The formations of Cotton and 
Rees. or a combination of them, would be of world-wide occurrence, and sub- 
divisions could well express the characteristic algal groups of the geographical 


The occurrence of marine algae m distinct sjoaes between and often below 
tide levels is a distinctive feature of rocky coasts, though more prominent where 
(he tidal range is large. The tidal range around Kangaroo Island is small 
(between 2 and 4£ feet), hut zonatlon is always present and often marked. 

In de1inutmg\he zones around Kangaroo Island absence of accurate tidal 
data 19 a limiting factor. Until such data become available, the position of the 
zone* oi algae in relation to tide levels can be given only approximately, and 
what appear to be critical levels only from subjective observations. Thus the. two 
main littoral associations of rocky roasts are referred to as being m the upper 
littoral" and "lower littoral/' but the relation of these to tide levels cannot be 
given. The upper littoral rone of blue-green algae probably does not extend to 
high water mark of spring tides, except when influenced by splash effects. 

A< Ion* as the occurrence of algae and animals on the shore is referred to 
as "zonaticu" -a word which is far too well established to he dropped— there 


seems no justification for replacing the term °zone M by "belt/' as is advocarcd 
by Chapman. 

It has been passible to relate the algal zones to measurements of tide levels 
only in the American River inlet. Here the stationary low water level of dodge 
Udes appears 10 be of most importance, marking the separation of ihe Honnosint 
zone from a zone of red algae (Hypnea- Centrocerax - Spyndia) which is nearlv 
always covered. This level is very little higher than the low water level of neap 
tides (sec graphs, fig, 2). On the south coast the low water mark of neap or 
dodge tides in summer appears to correspond closely with the surface level of the 
Mat rock-platforms, and this marks a distinctive change in the algal flora. The 
higher mean sea level in winter may cause an elevation of the littoral flora, bUI 
the lower summer level will be the limiting factor at least for the more perma- 
nent algae. 

The littoral zone is therefore considered as ranging from the stationary low 
of dodge tide*, or the low water mark o[ ne*p tides to the upper limit of the 
algal vegetation. Accurate fixation of this level will have to await detailed tidal 

The term "supralittoral" is often applied to the zone ahove high water level 
of spring tides- Alternative names are the •'splash" or "spray" zones. Cotton 
(1912) has given good reasons for rejecting this term, and investigations around 
Kangaroo island support the view that algal vegetation above actual high water 
level is simply an upward extension of the upper littoral algae under the influence 
of shade and wave-splash. One exception to this lies in the occurrence of 
Prostola during winter at Pennington Ray and on Shag Rock in Pelican Lagoon, 
well above the area splashed by waves. This alga is subject to fine blown S prav\ 
but is as much terrestrial as marine. In both localities it occurs only where penguin 
and shag excrement is present. The lichen IJchina may occur in 'small patches in 
and above ihe splash area, and the mollusc Mehmphe unif^ciata extends rnanv 
feet above b»gh water mark. Apart from these associations the term supralmoral 
is of little use in describing the algal ecology of Kangaroo Island. 

Below the littoral is the sublittoral, which extends down to the limit of algal 
vegetation. The upper limit of the sublittoral. particularly on exposed rocky 
coasts, bears a distinctive algal flora, and this area, between low water mark of 
neap tides and extreme low water of spring tides, has been termed the "sublittoral 
fringe" by Stephenson (1939). The sublittoral fringe on rough coasts is exposed 
during the suck back between waves at low tide, and the short but frequctit 
periods of exposure to air are probably of importance in determining the algnl 
flora present.. On the south and west coasts of the island this zone is dominated 
by Lystophora intermedia, which is strictly confined to the region exposed 
between waves at low tide. On calmer rocky coasts other species of Cystopftora 
are dominant, but these extend to 6 or more feet below low water. At American 
River also the flora just below low water is not so distinctive, and is better 
referred to as "upper sublittoral/^ The sublittoral fringe must be regarded 
simpiy as a useful division of the sublittoral in certain areas, such as the rough 
coasts of Kangaroo Island. 

This account is of a general nature only. While it is derived from the study 
of localities illustrating most of the coast and appears at the present stage well 
founded, it makes no pretensions to be final, and modification may be necessary 
as other coastal areas are visited. Descriptions of the typical localities will be 
given in later papers, 


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Fig. 3 shows the basic zonation at the localities studied. Two main reports 
are clearly defined: 

(}) The American River tidal inlet where species of Cyxtophora (or other 
large brown aJgac) are almost completely absent from the upper suh- 
lutoral zone. Hormosira banksii Dene., Getidium pusi/lum (Stackh.) 
f.ejol and Bostrychia simplichiu-itfa Harv. dominate the basic littoral 
zones from lower to upper littoral. 

i2} F^t rcSt ° f the coast ol the islancI >vhere s P e cies of Cyslophora (or 
Ecklonm radmta (Turn.) J. Ag.) arc dominant in the upper sub- 
iutoral zone, Coralline-mat and blue-green algae here form the two 
characteristic zones of the littoral. 
Hence depending on die presence or absence of species of Cvstophont t the 
coast may be divided into two formations named from their characteristic habitat. 
!'!? t ba ™ or Sandy-mud (Flat) Formation" (American River inlet) and the 
Rocky Coast Formation" (see fig. 4). The naming of these formations on ihe 
habitat has been discussed under "Terminology/' 

The chief differences between the two formations are: 
1.1 I The difference in species composition. Species common to both forma- 
tions are rare, and if commoh they usually dhTer greatly iri relative 
abundance. The Gelid imn pusilhun association, well developed and pro- 
minent in Pelican Lagoon, is present, but poorly developed at Pennington 
Bay, and fragments may be found elsewhere along the coast. Hormosira 
banksii forms well-developed associations in both formations, but ihe 
ecological forms in each are very distinct (see pk B, f\g. 4, and pi. E, 
fig. 4). In number of species the Rocky Coast Formation is much richer 
than the Sand and Sandy-mud Flat Formation, while the size of the 
algae is usually greater in the former. 

(2) Methods of attachment. A wide, expanded, holdfast disc is charac- 
1 eristic of all the larger algae in the Rocky Coast Formation. In rough 
places this disc is extremely strong. Other methods of attachment are 
found in the littoral zone; zns. } ramifying and densely matted attaching 
filaments of the coralline mat association, and the gelatinous adhesive 
ihalli of the blue-green association of the tipper littoral. 

in the American River inlet the expanded holdfast disc is rarely 
found, and the adhesive ihalli of blue-green algae are almost completely 
absent from the upper litioral. The chief mode oE attachment is by 
rrnzoidal filaments (a notable exception heing Hormosira). Attach- 
ment of filaments by means of a basal cell or cells occurs in both 

(,l) The growth substratum. On rocky coasts the vast majority of algae 
grow on rock, while a few occur as epiphytes where the growth is dense. 
Many epiphytes can grow equally well on rock or on other algae, but 
a few are limited to particular hosts (e.g., Notheia on Hormosira). ' 

At American River most of the tidal flats are colonised by the 
marine Angiosperms Posidonia australis Hook, and Zostera ■muelteri 
Irmisch, and these, hear a profusion of epiphytic algae. Apart from 
Bostrychia, Gelidium and Porphyra, which occur on rock in the upper 
littoral, and to some extent Hormosira, all others are cither epiphytes 
or ^row on shells and small stones in the sand and mud. No macro- 
scopic algae are able to grow directly in or on the sand or mud. although 
-hey may often be partly buried in mud; growth in sand ur mud is. how- 
ever, characteristic of the marine Angiosperms 


In discussing the Rocky Coast Formation, no account is taken of stretches 
of sandy beach between rocky sections of the coast. Rees considered such areas 
as a separate sub formation, but devoid of algae. No macroscopic forms are found 
on such beaches, and wherever rocks occur the typical algae of the Rocky Coast 
Formation are found. 

Within the Rocky Coast Formation conditions of roughness vary from very 
rough tti moderately calm, and the coast may be divided on the presence or absence 
of Cystophora intermedia. This brown alga is found only in conditions of fairly 
strong to very strong wave action, and never occurs on calm coasts. It grows best 
under the constant action of heavy breakers, and has by far the strongest thallus 
for its thickness of any Kangaroo Island alga. Cystophora intermedia is dominant 
in the sublittorai fringe zone from Cape Willoughhy along the south and west 
coasts, and along the north coast to between Western River and Middle. River, 
where it is replaced by other species of Cysiophora, 

a ,>„«. 



Sin J tsnd 5jni^u-rr'u4 Formation. 

Fig. 4 

The Algal Formations and Suuformatinns aramd the Kangaroo Island coast, 

Areas in the Bay of Shoals and Western Cove not yet studied are left unclassified 

Only the locialitles actually examined are shown on the map. 

On the north coast between Cape. Willoughbv and Middle River (excepting 
the American River inlet) other species of Cystophora (C\ subfarcinata (Mert.) 
J. Ag., C. siliquosa J. Ag., and others) or Ecklonia radiata, and in some places 
the red alga Acrotylus atistraiis J. Ag., are dominant in the tipper sublittorai zone. 
These species require constant water movement, but sudden rough weather will 
remove many plants from the rock. The outer edge of the Pennington Bay rock 
platforms is very rough, with calmer conditions nearer in and at the rear of 
the reefs, where Cystophora sitbfanmata and C. siliquosa are very common. Along 
the whole south coast, however, C. intermedia is dominant in the sublittorai fringe, 
and although fairly common, the species characteristic of more sheltered coasts 
occur only where conditions are locally less rough. 

The Rocky Coast Formation is therefore divided as follows (see fig. 4) : 


(1) The Exposed Rocky Coast Subformation; from Cape Willoughby 
along the south, west and north coasts to between Western River and 
Middle River. The area is characterised by the presence of Cystophora 
intermedia in the sublittoral fringe. 

(2) The Sheltered Rocky Coast Subformation; found along the north 
coast between Cape Willoughby and Middle River, excluding the Ameri- 
can River inlet. Characterised" by other species of Cystophora, Ecklonia 
radiata, and in some areas Acrotylus australis, in the upper sublittoral, 
Sargassum spp. may also occur in some areas. 

Cotton found it necessary to divide his Rocky Shore Formation into Exposed 
and Sheltered Series. This appears to correspond closely as far as environment 
and status go with the two sub formations of the Rocky Coast Formation on 
Kangaroo Island. 

Although referred to as "sheltered," the degree of shelter in this sub- 
formation is very much less than in the Sand and Sandy-mud Formation. The 
latter is developed mainly in almost land-locked areas, whereas the Rocky Coast 
Formation is always found on open coasts. 

Within the exposed Rocky Coast Formation two distinctive types of habitat 
occur, dependent on the geology of the coast (see "Coastal Geology," under 
"Environment"). These are the horizontal sand rock reefs, actually wave cut 
platforms, occurring along much of the south coast (pi. ix, fig. 3), and the steeply 
sloping rocky areas occupying the rest of the. coast (pi. xii, fig. 1), A brief 
description of the main associations in these two areas is given below, but detailed 
reports will be left till later papers. 

(it) The Pennington Bay Rock Platforms 

The type of horizontal wave-cut platform (pi ix, fig. 3) found at Penning- 
ton Bay occurs along much of the south coast of Kangaroo Island (see fig. 1)- 
The reefs which have been studied in detail at Pennington Bay are probably 
representative of this type of algal habitat, and a detailed account will be given 
in another paper. The following are the main associations found in the Penning- 
ton Bay area : 

(1) The Littoral Zo^e. 

Rear Littokal Associations — These occur on the vertical or sloping 
rock backing the reefs, usually at a higher elevation that the reef itself. The 
associations are exposed at low tide, but washed or splashed continuously 
at medium and high tides. 

1. Rhntkvriu firma association (pi. x, fig. 2). This alga favours areas where 
wave splash is moderate or else there is constantly running water. For 
further notes see Womersley (1946 a). 

2. Symploca hydnoides association: forming scattered patches in shaded 
hollows of vertical or sloping rocks. 

3. Gelidium pusillum association: common, but usually poorly developed. 

4. Enteromorpha association: forming bright green, usually pure areas on 
sloping well-washed rock. 

5. Ectocarpus confervoides and Pyhdella seasonal associations. These form 
brown mats and tufts on well-washed sloping rock, Ectocarpus occurring 
during winter and Pylaiella mainly in summer. 


Littoral Associations (on the flat reef surface). 

6. The Cystophora complex. Four species of Cystopkora, C. suhfardnata, 
C. siliquosa, C, nvifera (Ag.) J. Ag. and C\ broumii (Turn.) J. Ag., 
together with Sargassum tnuriculatum J. Ag.. form a complex of associa- 
tions on the flat reef surface, always where they arc submerged. Each 
species may form a pure association or occur mixed with one or more of 
the others, depending on the depth of water at low tide (pL x, fig. I), 
This complex covers the larger part of the reef surface. 

7. Hornwsira banksii association. This is a well-marked association on 
slightly higher and therefore more exposed parts of the flat reef surface. 
On higher areas it is pure and dense (pi. ix, fig. 4, and pi. x, fig. 4) ; 
in other places it may become mixed with species of Cystophora. Noihcia 
anofnaia Bail, et Harv. is always found growing from the conceptacles, 

8. Cystophyllum muricainm association. A pure area of this alga, occurs in 
well-washed, relatively calm, and rather sandy parts of the main reef. 

9. Laurevcia hetrroclada association. This occurs in fairly rough places, 
where it forms a dense mat of stunted plants. 

10. Cystopkora - coralline association, A distinctive association found on the 
rougher parts of the reefs, consisting of Corallina cuzneri I^amour. on 
rock and Jania fastigiata Harv. on species of Cystophora; a dense and rich 

(2) The Surlittoral Fringe 

11. Cystophora intermedia association. This brown alga dominates the outer 
edges of the reefs in the roughest conditions. It is an extremely rich 
association of small, often stunted species (over 50 have been recorded 
from an area of a few square yards), completely covering- the rock 
(pi. ix, fig. 2). ' 

The sublittoral assemblage will not be dealt with here, but it is very similar 
to that listed for Vivonne Eay (see p. 244). The coast at Penningtou Bay is 
very rich in number of species; over an area of § mile more than 220 species 
have been recorded, and many more, no doubt, remain to be found. 

(b) Steeply Sloping Coasts 

Three main zones, each comprising one association and in certain localities 
others, occur in the intcrtidal areas of the more steeply sloping parts of the south 
;m(\ west coasts. Heavy wave splash or regular passage of breakers up sloping 
rocks results in considerable upward extension of these zones. 

1. RivuLARrA - Isactts association of the upper littoral. 

This association is composed of Rhntlaria firma Womersley, R. atra Roth, 
and Isactts plana (Harv.) Thuret, all forming scattered, dark blue-green gela- 
tinous thalli on otherwise bare rock. In .some areas they are very well developed 
(especially R* firma), in other places they are almost absent. Degree of wave 
action is the determining factor, but the association is often poorly developed 
where least expected. 

2. Coralline-mat association. 

The lower littoral, between the blue-green algae and the sub-littoral fringe, 
usually consists of a dense mat of stunted Jania fastigiata and /or Corallina (pro- 
bably pi cwvieri) (pi. xi, fig. 4). This mat is 1-3 cm. in thickness, pinkish-white 
in colour, and forms a continuous covering on much of the rock. Where breakers 
run well up sloping rocks it may reach a height of 5 or 6 feet. At Vivonne Bay 


(on the south side of Ellen Point) t Dasyopsis ciamgera Womersley, and a small 
Start Laivrencia are prominent amongst the corallines. This jQnta-DQ-syopsis- 
Lourtntcia variant is probably general in many areas of the south and west coasts, 
but rather than being a distinct association it consist of the addition of the latter 
two algae to the basic coralline-mat association. 

3. Cvstophora intermedia ASSOCIATION of the sublittoral fringe. 

This brown alga forms a striking sublittoral fringe zone on sloping rocks 
of the south, west and north-western coasts of Kangaroo Island. The upper edge 
of the association is often very sharply limited, as shown in pi. xi, fig. 2, the zone 
appearing as a dark band stretching along vertical rock at Cape Willoughby. In 
situations where waves pass along the rock, rather than breaking against it, the 
coralline-mat and blue-green zones may be poorly developed, but the sharpness 
of the upper limit to Cyslophora intermedia can be seen from pi. x, fig. 3. Where 
waves break heavily on rocks the upper edge is less well defined, and the coralline- 
mat often merges with Cystophora intermedia (pi. xi, %- 4). 

The dark-brown pinnate fronds of Cystopfwra intermedia reach a length of 
40 to 45 em. The stems are extremely strong, and only very rarely are fronds 
ftmnrl cast up. A common epiphyte is Corynophhea cystoplwrae T, Ag. 

The Sublittoral. 

Study of the sublittoral flora is restricted to the algae cast up but not known 
to occur in the intertidal area. The following list includes the commonest forms 
of the sublittoral assemblage of the <outh coast, but comprises only a small 
fraction of the total. 

Oilokopiiyceae— Caulerpa harztcyi F. v. M.; C. obsnt/a Sonder; C. vesicuUfcra 
Harvey: Codiunt goleahtm 1. Ag. ; C\ nmndlhsitm Harvey; C. pomoides 
J- Ag. 

Pkaeofhyceae— Phloeocantofi spectabile Reinke ; Dictyota latifoHa J. Ag. ; 
Zmaria tumcriana f. Ag. ; Sporochnns scoparius Harvey; S. cowosns C Ag.; 
Bdhtia eriophoriim Harv.; EnryotJiolia cliftoni Harvey; Peritl\alin inenms 
(R. Br.) |. Ag ; Eckloma radiola (Turn. 1 ) J. Ag,; ScytathaHa dorywtpa 
(Turn.) GrcvT; Sierococcus axillaris Greville; Scaherh agardii Grev. T 
Myrwdesma qucrcifol'um (Bury) J, Ag.; Carpoglnssam confluem (K. Kr.) 
Kfitz.; Cvstophora mow'lifera |. Ag. ; C dumasa J. Ag.; C, retorta (Mert.) 
J. Ag.; C. rawvtosa Harv. : C, platylobium (Mert.) J. Ag. , C\ spnrtuudcs 
(Turn.) J. Ag.; C. siliquosa J. Ag. ; C punindata (Turn.) J. Ag.; Sargas- 
stiw varians Sonder; .9.. sondrri ). Ag. ; S. tricfwphylhim J. Ag. ; S. enstatum 
3 Ag. 

KhoI'OPhvceae— Asparagopsis anmtta Harvey; Mychodea cuwpressa Harvey; 
Hypnea episcopatis II. & H.; Dclisea vh-gans C Ag. ; Phaceloearpus Iabdiar- 
dieri J, Ag.; Plocamhm nidipcum (ITarv.) J. Ag. ; P. preissianum Sonder; 
P. costotum (J. Ag.) H. &. II.; Hymcnochdia potymnrphn (Harv.) J. Ag.; 
Jtttithanmwti wucmnatum (J. Ag.) Rte Toni ; Momhspora olongata- (Harv.) 
Dc Toni ; Ct>ramium pitberulutu Sonder ; iMstothalia furmosa (Harv.) 
Dc Toni ; Spongochvnnn sp.; Spyridia opposUa Harv. ; Sarcomenw dosyatdes 
Harv. ; Nitophytluw. curdieanum Harv. ; Amansia pinvatifida Harvey ; 
Li-normandia spectabilis Sonder; Osmundaria prolifera Lamour. ; Tharctia 
iuicrafolia Dene. 

Other Communities of Steeply Sloping Coasts 
An Entcromorpha association occurs on ruck well above normal wave-splash 
at Cape Coudie. It is "dependent on the presence of fresh water percolating 


through the tipper limestone stratum and running clown over the harder ancient 
rocks forming the base of (he. cliffy. Hie species has not been determined, but ir 
occurs in dense, pure, masse? on otherwise bare rock. Enteroumrpha associations 
dependent on the presence of fresh water have been recorded by numerous other 
authors (see Cotton 1912), 

A Splarlimdnim rugosum association is found in the upper littoral at Cape 
Willoughby, usually at a higher level than the hliic-grccn zone. The Cape is 
composed of granite boulders, and where waves break heavily, leaving the rock 
exposed between waves, Splachmdium forms a pure association of short, tufted 
plants (see pl v xi, fig. 3). 

At Vivonnc Bay, on gneissic rock, Sphichnidiwm rtujosuin (L.) Grev. is often 
common on wave-splashed rock, but may merge with Rivnlaria firina. Hdvw- 
thora tumens J. Ag. and Pidyslphonia dasyoidea Zan. are also characteristic of 
this region during January. 

On other types of rock Splachnidiuni h very rare. If further studies show 
that it is restricted to granite or gneissic rock, this will he one of the very tew 
cases known from Kangaroo Island of the type of rock influencing algal 

A marked feature of the south side of fallen Point, Vivonne Bay, :"s the 
occurrence of at least five species of litholharnnia. Elsewhere on the island they 
are rare. Two distinct species (generic determination has not yet been possible") 
form pure but localised communities in the littoral zone in what are apparently 
rather specialised habitats. They grow as crustose thalli forming small irregular 

The coast at Ellen Point consists oT f ossiferous calcareous limestone over- 
lying the hard gneissic base. Weaving back of the softer limestone hat resulted 
in many rock pools, from very smad to over 20 yards across, being left 
in the harder base (pi. xi, fig I), Most of these pools are subject to 
wave influx only al high tide, and during summer their wafer temperature is con- 
siderably higher than that of the sea ( up to 28^ C when *ea temperature is 
18° C ). 

Some of these pools bear distinctive algal communities; others, where con- 
ditions are apparently too severe, are devoid of growth. 

One pool, shown in the foreground in pi. xi, fig, 1, contains a Lamrncia — 
lithothamnion community. The lithothamnion forms scattered, irregular pinkish 
masses (to 10 cm, across and 3 cm. thick), while Laurcncia Iwterachido Harv. 
grows on the rock or the lithothamnion and is heavily epiphytiscd bv Ccramium 
miniattnu Suhr. and Polysiphonia abscissa Harv. 

In the rear pool of the two shown in pi. xi, rig, 1, the end shaded by the cliff 
hears a community of red algae which are normally suMiuoral forms. Dictymenut 
iridens Grev. and Bamctia sp. are the commonest, while in another shaded area 
of the same poo! Lyncjhya majuscitla (Dillw.) Hurv. forms a pure community in 
January. The effect of continual shade is evident in both cases. 

In another pool, at a lower level and .subject to wave influx except, at low 
tide, species of Cystnphora are dominant. One corner, however, is shaded by 
overhanging rock, and here Ecklonia rodiala, Srytothotia dorvcarpa, Myrmlr.umt 
Intifoha Harv. var. durhtscuto J. Ag. (with epiphytic Sphacvhiria iributoides 
Meneg(i.),and Gelidhtm frustrate J. Ag. are prominent. All Ihese arc normally 
upper sublittoral forms. 

Tn £-omc of the pools minor communities of Bryopsis plmnosa (Huds.) 
C. Ag., Hryop.sis bncuHfcra- J. Ag.; Derbeda sp., two h'thothamnia, and 
two species of coral occur. This assemblage shows more relationship to the flora 
of tropical waters, and is almost certainty due to the higher temperatures main- 
tained in these pools during summer 


To deal adequately with the complex nature of the littoral zone at Ellen 
Point requires detailed mapping of the greatly dissected coastline. The variation 
in minor habitats is almost without limit, and similar complex areas probably 
occur along other parts of the south coast. However, the basic zonation of blue- 
green, coralline-mat and Cystophora intermedia zones is found on all rocks 
directly exposed to the sea. 


Wave action on the coast included under this subformation is from moderate 
to slight (see pi. xii, % 2 and 4). In fine weather waves gently lap the shore, 
while breaJcers a few feet high occur in rough weather. Some degree of water 
movement is always present, whereas in the American River inlet conditions are 
more often than not a dead calm on the tidal flats. 

Littoral zonation is basically similar to that on exposed coasts, comprising 
Wue-green and coralline-mat associations. 

1 Thk Upper Littoral Zone of Bltje-grken Auiae. 

On the eastern end of the island, and at Middle River, Rkmlaria firm* is 
dominant, accompanied by Rivularia atra, Isactis plana and sometimes Sympbca 
hydnoides Kutz. In calmer areas R. firma disappears and R. atra and Isactis 
become dominant From Middle River to Stokes Bay (and probably further eastl 
Brachytrichia quoyi (Ag.) B. & R is prominent in January. 

In some places where wave-splash is absent this blue-green zone may be very 
inconspicuous. The gelatinous thalli occur scattered singly or in patches on other- 
wise bare rock. No other algae normally occur in this area of the upper littoral. 
Blue-green algae are absent at Rocky Point, where the sxibstrate is consolidated 
sand rock and the littoral zonation is closely allied to that found in Pelican 

A community of Ncmalion helminthoides (Velley ) Batt. occurs on rocks on 
the east side of the beach at Middle River, in the mid-littoral, while on the west 
side of the beach Ctadosiphon (Hum (Harv.) Kyiiti is common below the blue- 
green zone. 


This is usually well developed in the lower littoral, often forming a closed 
community with a well-defined upper edge (see pi. xii,. fig. 4, at Emu Bay) . Jama 
fastiguita and fragments of Corallina are the main constituents, but Dasyopsis 
clavigcra and the Laur^nda of the south coast association are absent. Gelidwn 
jrusillum Is often, in calmer localities, an integral part of the mat, while other 
species commonly present are: Wrangelia plumosa Ifarv., Rachydictyon pam- 
lulahtm J. Ag., Zonatia- turnerimui J. Ag., Am-phima charoides Lamour, 
l.aurenda hcteroclada Harv. s Ceramium miniatum and Polysiphonia spp. 

At Rocky Point Gelidium piisillum has become completely dominant, with 
only fragments of coralline left amongst the Hormosira association at a lower 
level. Above this is a zone of Bostrychia, similar but often better defined than in 
Pelican Lagoon. 

3. The Upper Subl.ittoral Zone. 

This region, on sheltered rocky coasts, is characterised by the dominance of 
fairly large brown algae, forming several associations in different localities. 

Cvstophora association. 

Three species, C. subfdrcinata, C. potyrystidta Arcsch. f C. STliquosa, and to 
a lesser extent C. spartioi-des, extend from low water mark to a depth oi 6 or 
8 feet. They may reach a length of H meties, and at low tide in some localities 


the fronds float at or below water surface, giving the zone a distinctive 

C. snbfarcinata is found throughout the subformation, but in calmer areas 
{e.g., Rocky Point) forms numerous vesicles and is more branched. C siliquosa 
and C. spartioides are restricted to slightly rougher parts, while C. polycystidia 
becomes dominant in calmer regions. In shallow water and locally calm places 
Cysfophyllum mnricatum forms a distinct community* 

Although G polycystidia is characteristic of the north coast of Kangaroo 
Island, it does occur in locally sheltered places on the south coast. 

On the east side of Ballast Head (north of American River) San-jassum 3p. 
dominates the upper sublittoral. 

Eckhnia radiatu association. 

At Hog Bay and Rocky Point, where wave action is slight, Ecfzlonia r&diata 
forms a distinct sublittoral fringe, accompanied by some C. snbforcinato. Under 
locally suitable conditions elsewhere around the island it may be found; at Cape 
Coudie a protected channel bears a dense fringe of Ecklonia, 

Acrotylus australis association. 

At Middle River and the east side of Cape Willoughby dense and pure 
patches of the red alga Acrotylus australis occur in the upper sublittoral. The 
dark brown dichotpmous fronds, from 10 to 20 cm. high, completely cover the 
rock, forming a distinct association. At both localities conditions of wave action 
are very similar, and the general algal ecology is almost identical. Caulerpa 
brownn Endl. often forms dense bright green mats within a few feet of low 
water mark* 

The following species are commonly cast up within the Sheltered Rocky 
Coast Subformation ; Codium spongiosum Harv., C. pomoides, Cladostephus 
vertieillatits ( Light f.) Ag., Halopteris pseudospicaia Sauv., Saryassum sp. (small 
stunted plants), Cystophora bolryocysth; Sonder, C. grevillei (Ag.) J. Ag., 
C. moiiUifera J. Ag., Amphiroa charoides, Dictymenia harveyam Sonder, 
Lattrcncia sp. 

In Kastern Cove, and probably in Western Cove, at least four distinct asso- 
ciations occur in deep water, 

(1) Posidonia australis, known a? the "sea grass" or "tape weed," forms 
extensive meadows on a sandy bottom in from 1 to 6 or 7 fathoms of 

(2) Scaheria agardhii Grev. occurs on a rocky bottom in from -J to 3 or even 
5 iathoms below low water. 

(3) Cystophora monttifera occurs in from 1 to 7 fathoms, 

(4) Chimcanthia arbor ca (Ilarv.) Falk. forms dense masses, especially in 
winter, in 1 to 2-J fathoms. 

The shore in the Bay of Shoals and Western Cove is sandy and muddy, with 
few rocky areas. Tidal flats, however, are not formed to any extent. This type 
of hahitat is intermediate between the Sheltered Rocky Coast Subformation and 
the Sand or Sandy-mud Formation, with closer affinities to the latter, Until the 
area has been more thoroughly investigated, no classification will be attempted. 

Along the shore of the Bay of Shoals Zostera muelleri is common, and 
amongst it, on old shells, occurs the green alga Acetahularia penicuius R. Bf. 
(probably a winter form). In deeper water Posidonia australis is dominant. 

Rock Pool Associations, 

At Western River and Middle River rock pools are a feature of the coast. 
They are mostly small, from 1 to 10 feet across and to 2 or 3 feet deep. During 
summer the temperature in smaller pools (containing Enteromorpha) reaches 


35° C. The conditions in any pool depend on its size, height above sea. level, and 
general situation; the environment of smaller pools during summer is extreme in 
both temperature and salinity conditions. Two types of pools occur: 

(1) Entckomorfh a Pool Association. 

This association occurs in the smaller and higher pools where conditions 
are extreme and v^ry variable. E, lingidaia J. Ag. and H. intesthialis 
(L.) Link, form a dense fringe around the edge. In summer exposure 
on the water surface often kills and bleaches the upper plants. The 
toleration these algae have for high temperatures is shown by their 
active oxygen liberation under temperatures of 30-35° C. 

(2) Hormosira - Cystophora Poor. Association. 

In larger and lower pools, where waves enter more frequently and 
temperatures are therefore lower, Hormosira hanksii forms a dense 
fringe around the edge, at or just below water level (see pi. xii, fig. 3). 
while Cystophora suhfweinata, C polycystidia, C, siliqunsaj. C. brmvmi, 
and often Cystophyllum murkatum and Sargassuw sp. grow on Ijifc 
lower sides and bottom. 

In many pools along the north coast, particularly those with a sandy bottom, 
the only algal growth consists of small mats of Grlkfium pusiUum and fragment* 
of CoroUina and Jania. 


The American "River tidal inlet comprises several large lagoons with wide 
tidal fiats and a central channel, opening into Eastern Cove through a narrow 
■neck. Conditions are very calm, particularly on the tidal fiats where large beds 
of Fosidonta and other weed tend to minimise wave action. 

Tidal range is just over 4 feet, decreasing only 2" or 3" from American 
River to Pelican Lagoon (see fig. 2). The fast scouring currents during spring 
tides, together with the sandy bottom, prevent algal growth in the channel proper. 
The temperature range on ihe flats is large, for at low tide less than a foot, and 
often only I" or 2" of water covers. t]ie algae. Winter temperatures reach as 
low as 10 D C, summer up to 32° C on the flats, 

In Pelican Lagoon the tidal flats usually extend out from low cliffs of cal- 
careous sand-rock (similar to the coast at Pennington Bay) (pi. xiii, rig 2), hut 
sandy beaches are frequent, especially between Muston and American River jetty. 
South of Muston samphire swamps cover several miles of the shore. 

An important characteristic of this formation is the large quantity of move- 
able sand and mud. The fast tidal currents carry suspended mud, which algae 
on the flats must be able to tolerate. The characteristic colour of Rhodophvccac 
at American River is a dirty brown, very different from the red of clean water 
forms at Pennington Ray. 

The basic zonation in Pelican Lagoon is shown in fig. 5. Where a sandy 
beach occurs, Hormosira or Zoslcra comprise the upper zone. Over most of the 
flats the area colonised by Hormosira, Zostcra and Posidonia is much greater 
than shown in the figure, but the sequence of infest and their positions in relation 
to tide levels applies generally. Microscopic algae, particularly diatoms, arc 
usually present as epiphytes on the larger algae, but identification has not been 

On Shag Rock (a small island at the entrance of Pelican Lagoon), and 
probably elsewhere where shag colonies occur, a filamentous form of Prasiola 
covers rocks affected by the bird droppings. This appears to be a winter associa- 
tion only, occurring several feet above high tide level. 


Littoral Associations 
1. bostrycitia and gttudium associations. 

These two associations will be considered together, as they occur in very 
similar habitats and sometimes become mixed. At Rocky Point, however, each 
association forms a distinct zone } She Gciidium below the Bostrychia. Roth algae 
form dark reddish-brown dense mats, up to 1 cm, thick, covering the rock from 
about mid to just below bigh water level of spring tides. They are restricted to 
shaded areas of rock (see pi. xiii, fig. 2, and fig. 5), With continual shade, the 
mats retain sufficient water during the periods of exposure. Rock not shaded in 
this zone is usually bare of macroscopic algae. 

Fig- 5 

Typical algal venation on Pig Island, Pelican Lagoon. The relation of each zone 
to tide levels is approximately true; but in many areas* the tidal flats are much 
v.^cit.r and the Hormosird, Zostcra \\\\(\ Poshionna assentations occupy far greater 
ureas than indicated. H.W.S.. tsiyh water springs; \T..\V_S,, low water springs; 
Il.W.N., high water near'-*: S.W.O., stationary water dodges. 

As a general rule, Gclidiitm pusiUum covers lower and more exposed rock 
than Bostrychia {B. $im[>lkhiscula, with some B. mixta H. & H.)- The roof and 
sides of small wave-cut caves are the most suitable habitat for Bostrychia. 

During winter a community of Porphyra unibiUcalis (L.) J. Ag. occurs in 
the lower Gelidhtm zone, but on more exposed rocks. The red-purpie fronds 
reach a length of only 8 to 12 cm., and lie as flat sheets on I he rock al low tide. 

From the. base of the low cliffs to the start of the Hnrmosira association (a 
distance of up to 20 yards) is a zone bare of macroscopic algal growth. The area 
is left quite dry and fully exposed to the sun at low tide. Such conditions are too 
severe for any alga, but the molluscs Bemhicium melanostoma and Modiolus 
areolatus are common. 


2. Hormosira Association. 

Hormosira banksii forms an extensive and conspicuous association in the 
low littoral. From about 1 foot above the dodge low to an inch or two below. 
Each plant grows either on the rock, or, if the substrate is sandy or muddy, oil 
the partially buried bivalve Brachyadvntcs erosus. The spherical "water contain- 
ing receptacles of Hormosira enable it to grow where it is always exposed at low 
tide (pi. xin\ fig. 4). 

The form of Hormosira banksii at American River is labillnrdicri Harv. 
Tt differs from the south coast form in being more branched, and having larger 
and more spherical vesicles. In both localities it grows only where it is exposed 
for a period each day; this exposure must in some way be essential to tbe growih 
of the alga. 

During winter Echicarpus confervoides (Roth.) Le Jol, is. a common 
cpIpTiytc on Hormosira at American River. 

3. Ulva Association, 

Ulva tacHtca is always prominent at about low tide level, in the lower 
Hormosira and upper Hypnea - Centroceras - Spyridia associations. On the flats 
north of the American River jetty it forms a green band along the shore, super- 
imposed on the Zostcra association. In other areas it frequently becomes dominant 
to form a distinct association. 

Forms of Entcromorpha prolifera J. Ag., E, dathrata Roth, and E. bulhosa 
Kiuz, are common in the lower littoral and upper sublittural, sometimes forming 

sublittouar. associations 

4. Hypnea - Centroceras - Svyridia Association. 

This association extends from about low dodge level to 1 foot or slightly 
more below. The dominant algae arc brown-eoloiired Rhodophyccae, and where 
dense give the zone an even brown appearance Hypnea muse if or mis ( Wulf.) 
Lamour,, Centroceras davidatuvt Ag. and Spyridia biannulata J. Ag. occur in 
varying but often about equal proportions. They are all slender forms, rarely 
more than 25 cm. long, and are just exposed at very low tides. Tn some places 
Gracilaria confervoides (L.) Grev. is common; on muddy patches around the 
small islands in Pelican Lagoon it forms dense but localised communities. The 
base of each plant is buried in mud but actually attached to a mollusc. 

Other algae occur in the lower part of this association, and in somewhat 
deeper water. The distinctive feature, however, is the virtual absence of any 
species of Cystophora or other large brown alga. (The one exception is Sargassmu 
(ufur!){(\ which occasionally grows on rock on the sides of the channel where there 
is more water movement, but rarely in the upper sublittoral.) The commonest 
species are: Caidcrpa remotifolia Sonder; C". simpliciascula; Codutm nmelU-ri 
Jvutz. r a lithothaumion forming spherical nodules to 5 cm. across; Corallina sp. 
(hcmi:rpherical tufts 20 cm. across), and more rarely Cystophyllmn muricaitim, 


Zoslera muellcri. forms a pure and dense association over large areas of the 
tidal flata (pi. xiii, fig. 3), from low water level to 2 feet below, in some 
places to as much as 6 teet below. This anglospcrm spreads almost entirely bv 
means of runners which anchor the plant in the mud and produce narrow leaves 
(to 25 cm. long). It seems to prefer a tnu«ldy substratum. Epiphytes on 
the leaves are common (pi. xiii, tig. 3), especially Centroceras clavulatum, 


Cladophom ceralina Kxnz., Ceramiuw sp., and Rivularia polyotis (Ag\) 
H, & ] 1 '. Amongst the Zostera, on small sandy patches, Chondria dasyphila 
(Grtiv.) C. Ag., Spyridia bknmufata, Polysiphamu patcrsonis Sonder and 
GracWarui conferva ides occur. Very rarely plants of Cystophytlum muri- 
cutitm and Cystophont ccphalornilhus (Lab.) J-Ag. maybe found. 


Pozidonia aust-ralis colonises deeper parts of the tidal flats and the channel 
edges, from 1 lo 10 feet below extreme low water, with occasional plants to 
,14 feer, Jn Eastern Cove it extends to about 7 fathoms. The long strap-like 
leaves have a distinctive appearance as tliey just reach the water surface on the 
flats at low tide. 

The association is dense and pure, but the rough leaves hear a wealth of 
epiphytes. On small pieces of leaf 10 species are nfu:u present. The most 
important ones are: Rwularia polyotis, Ectoctwp-us sp.. Asp QtO coccus bullosas 
Lamour,, Colpomenia sinuosa (Roth.) I). & Sol,, Jan'ui mknirthradw Lamour., 
Caitmceras elavulatuin, Ceramium paberulum, Spyridht bianmtlata, Polysiphonia 
sitcculenta Harv., P. fuscescens Harv., P. davyae Reinb. Many of these attain 
their maximum development during winter months. 


Another marine angiosperm, Hulophila ovalis (R. Br.) Hook, forms dense 
patches between 2 and 12 feel below low water. The ovate leaves, produced from 
runners, reach a height of 15 cm., but epiphytic growth on them is much less than 
on. the Zosicra or Posulonia. 

The three marine angiosperms all spread largely by runners. This ensures for 
the most part a pure association, but in shallow water all three have been observed 
growing together. 


The author is indebted to Professor J. G. Wood for advice during the 
progress of this work and criticism of the manuscript. 

In much of the field work the author has been fortunate in having the 
assistance of members of the staff and senior students of the Botany School, 
University of Adelaide. Mr, S- J. F.dmonds, of the Zoology Department, has 
also been studying the animal communities of (he Kangaroo Island coasts, and 
most of the hydrological data were taken jointly by Mr. Edmonds and the author- 
The following have also rendered invaluable assistance in determinations of 
certain algal groups; Dr. Francis Drouet, Chicago Natural History Museum, 
U.S.A. (Myxophyceac, excluding Rivularia) ; Dr. H, K. Phmney, Chicago 
Natural Hislory Museum (C'ladophoraceae) ; Miss L. K, Moore, Holany Division, 
D- S. & L R. t New Zealand [Stypocaidaceac) ; Miss V. May, Fisheries Division, 
C. S. & 1. R-, Cronu!}a, N.SAV- (specie* mentioned in census and confirmation 
of others), The Directors of the Melbourne and Sydney National Herbaria have 
given most_ valuable assistance in the loan of named material. Mr. D. J. 
Rorhrford, of the Fisheries Division, C. S. & I. R., Cromdla, has carried out 
phosphate and nitrate analyses of sea water samples from Kangaroo Island. To 
all these my thanks are extended. 

Baldwin, J. G-, and Chockkh, R JL. 1941 "Soils and Vegetation of Portion of 
Kangaroo Island, South Australia,/'" Trans Ew, Soc. S. Aust t 65, 

CiiM'MAK, R. W. 1924 "The Tides of the South Australian C^tsL" Aust 
Ass r for Advancement of Science,. 17th meeting, Adelaide 


Chapman-,. R, W. 1938 "The Tides of Australia/' Official Year Book of the 
Commonwealth of Australia, No, 31, 972 984 

Chapman, V. J. 1946 "Marine Algal Ecology." Botanical Review, 12, 

Clexaxd, J. P>., and Ruack, J. M. 1941 "An Enumeration of the Vascular 
Plants of Kangaroo Island. Additions and Corrections/' Trans. Roy. 
Soc. S. Aust., 65, 244-248 

Cotton, A. D. 1912 ''Marine Algae/' Clare Island Survey, Proc. Royal 
Irish Academy. 31, tit. 15, 1-178 

Crocker, R r L., and Wooo, J. G. 1947 '"Some Historical Influences on the 
Development of the South Australian Vegetation Communities and their 
bearing on Concepts and Classification in Ecology/' Trans. Roy. Soc. 
S. Aust., 71, 91-136 

Harvkv, W. H. 1854 "Some Account of the Marine Botany of the Colony of 
Western Australia," Trans. Royal Irish Acad., 22, 525-566 

Hedley, C. 1915 "An Ecological Sketch of the Svdnev Beaches/ 1 Pme Roy. 
Soc. N.SAV., 49, 15-77 

Lucas, A. 1-J. S. 1929 "A Census of the Marine Algae of South Australia/' 
Trans. Roy. Soc. of S. Aust., 53, 45-53 

Lucas, A. II. S. 1935 "The Marine Algae of Lord Howe Island." Proc 
Linn. Soc. N.SAV,, 60, 194-232 

Madican, C T. 1928 "Preliminary Notes on New Evidence as to the Age of 
Formations on the North Coast of Kangaroo Island." Trans Roy Soc. 

5. Aust., 52, 210-216 

Pope, E. C. 1943 "Animal and Plant Communities of the Coastal Rock- 
platform at Long Reef T N.SAV." Proc. Linn. Soc. N.S.W., 68, 221-254 

Rees, T. K. 1935 "The Marine Algae of Lough Tne " Journal of Fxoloev 23 
69-133 w ' ' 

Stephenson, T. A. 1939 "The Constitution of the Imertidal Flora and Fauna 
of South Africa, Pt. T/' Linn. Soc. Lond., Journal, Zooloev 40, 

487-536 " hJ 

Svkrukcp, H. U., Johnson, M. W., and Fleming, R. H. 1942 "The Oceans. 
Their Physics., Chemistry and General Biology." New York 

Tanslev, A. G. 1940 "The British Islands and their Vegetation." Camhridge 
Univ. Press 

Tate. R. 1883 "The Botany of Kangaroo Island." Trans. Rov. Soc S Aust 

6, 116-171 " 

Wade, A. 1915 "The Supposed Oil-hearing Areas of South Australia/' Rtillc- 
tin No. 4, Geological Survey of South Australia 

Wood, J. Q 1930 "An Analysis of the Vegetation of Kangaroo Island and the 
Adjacent Peninsulas/' Trans. Roy Soc. S. AttsL, 54, 105-139 

Wokerslev KB. S. 1946a "Studies of the Marine Algae of Southern Aus- 
tralia, (fctl. The Genera Isactis and Rivularlar Trans. Roy. Soc. 
b. Aust., 70, 12/-136 

Womerslsy H. B. S. 1946b "Studies en the Marine Algae of Southern Aus- 

l£« m?i*J " CW Species ° f Das y°P x, s &0nl Kangaroo Island. 

laid., 137-144 

Trans. Rov, Sue. S. Aust.. 1047 

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ByL. C.P.Kerling 


During an investigation of pre-emergence rotting of peas at the Waite Institute it was evident that 
infections of seedlings by Pythium spp. were important. It was not known what species were 
concerned, and the writer undertook to investigate this problem during a brief sojourn in Adelaide. 



L. C. P. Kerltnc * 

Communicated by D. B, Adam 

[Read 11 September 19471 

During an investigation of pre-emergenee rotting of peas at the Waite 
Institute h was evident that infections of seedlings by Pytiiiuni Sf>#, were 
important. It was not known what species were concerned, and the writer under- 
took to invesiigate this problem daring a brief sojourn in Adelaide. 

Available for study were six isolations of Pythhtw from pea seedlings which 
had been grown in a red-brown rarlh at the Waite Institute, two from potatoes 
affected by "leak'' disease and one from a tomato seedling. The isolations had 
been made by Mr. N. T. Flenlje. 

The various isolates were grown on potato-dextrose, oatmeal, cornmeal and 
water agars, respectively, as used by Middleton (194-3), and on carrot agar as 
described bv Johann (1*928) and Schu'z (194-2). Since carroty agar, which has 
been auiocbved, was said to be unfavourable for oospore production (Sehulz), 
it was sterilised by steaming it twice at 95 R C. However, oospore reproduction 
was secured quite readily on the autoclaved media. The cultures studied were 
derived after a series of transfers from the edges of 24-26 hour old colonies, and 
each time the inoculum piece comained about half-a-dozen hyphal lips, Attempts 
were made io use single hyphal tip*, hut only in one case which is mentioned 
later was this successful. *The use of single zoospores was precluded because 
their production was never observed. In all cases germination took place by 
means of a germ tube. The observation of oogonial and antheridial characters 
was facilitated by using cultures grown on a drop of potato agar m a smalt 
moisture chamber made from a glass ring and cover slip, 

The diameters of the reproductive organs provide important criteria for 
differentiating species of Pythium, and it is desirable to measure large numbers 
of these structures to ascertain their range and mean value. It was found (hat 
diameters of oogonia and oospores were less variable than those of the sporangia, 
and that counts of 100-300 revealed thai they followed a normal frequency dis- 
tribution. It is appropriate then to describe the variability of means in terms of 
their standard errors calculated in the usual way. The sporangia, however, were 
much more variable in size and shape and only limits in size observed are 


It was evident that the isolates could be referred to Pythhtm ticbaryanttm 
Hesse or closely related species. 

The genus Pvthhim has been treated systematically at least sis times and 
has been the subject of several monographs. The isolates made locally have been 
referred to the species mentioned below and are reviewed m the light of recent 
descriptions and figures of Drechslcr (1946). Matthews (1931), Sidcris (1932), 
Van Luyk (1934, ajid especially, that of Middlelou (1943). 

* Laboratoriurn voor Mycologie an Aardappdonderzoek, Wagctiingcn, Holland, 
Traas. Roy, Snr. S. Atist., 71, (2), 1 December, 1947 


Fiy. 1 Pythium dvbaryanum Hesse 
A-D — Sporangia: E, Empty mono- 
clinous anthe-'.-jdhim; F, Diclinous an- 
thcridium; C, Oogonium penetrated by 
monoclinans and diclinoud antheridia. 

F'g. 2 Pylfiruni uJthnum Trow 

A - B — Sporangia : C— J : Oogonia and 

tospores earn with one monoclittotis 


Fig. 3 Pythvnn polymorphan Sideris 
A - D — Sporangia ; P, G, IT, Antheri- 
riiUrn after fertilization; I, Intercalary 
oogonium; J, E, O, Small oogonia with 
curved swollen antheridia; K, L, M v 
Young cog-onia with one anrlicridium. 
Antheridia) Halle typically curved; N, 
Two aiitlieridia to one oogonium, 

Fig. 4 Pylhhtm vrxans tic Bary 
A - C — Sporangia : D, Mycelial body ; 
J£, Germinating sporangium; F, Allamoid 
rmtheridium and young oogonium; G, H, 
J. O, Variations in antheridial shapes 
L, oogonium with two oospores; N, 
Aplcrotic oogonium; K, Germinating 
oospore with empty, antheridium; M, 
Germinating- sporangium. 


Pythium debar yanwtu Hesse 

Hyphae 2* 5-11 /a, usually 4-5 /<• in diameter; sporangia spherical to oval 
terminal or intercalary, sometimes in chains ranging From 9-41 fi m diameter and 
germinating by germ lubes; oogonia smooth, spherical, terminal, 14-29 #+ means 
20-7-23* 1 fi in diameter. Antheridia one or more per oogonium generally 
diclinous hut sometimes monoclinous when they arise some distance, 40-280 & 
below the oogonium. Antheridia stalked, crooked-necked with the terminal 
portion oblong spherical nv clavatc and 5-1 1 ;t in diameter, apex obtuse or narrow, 
tapering gradually »o the oogonial wall to which it is more or less applied. 
Oospores smooth, aplerotie, ll-25/t„ averaging 17-4-19'9/t in diameter, with a 
thick wall, Parasitic on Pisuui sativum and Solanum tuberosum (see lig. lj. 

The above description covers four of the isolates from peas and one from 
a potato affected by "leak." It agrees generally with those erven by other authors, 
and at this point only the sizes of oogonia and oospores described above cull for 
comment. If those quoted arc compared with those of various authorities [Butler 
(1913), Hrann (1925), Sidcris (1932), Van Luyk (1934), Middletoii (1943) L 
it is seen that the range of means we found is somewhat less than (hat described 
in the literature, Actually, statistical analysis reveals that the. means for both 
oogonia and oospores in our different isolates may be significantly different h'or 
example. the mean diameters of oogonia and oospores of isolate 5 from peas were 
20 '7 & ±0*09 and 17*4 /t^= 0*18 ^ respectively, whereas corresponding measure- 
ments lor isolate 2 (also referred to Pythium dfibaryanum), were 23'1/**=<M9 
and 19* n — 0*22 respectively. However, it is possible to arrange the isolates we 
classify as h'ylhium debavyanum in such U way that differences between succes- 
sive pairs are no| significant. The differences in means may indicate the existence 
of strains, but they offer little warranty frw specific segregation. 

Pythium ulthnum Trow. 

Hyphae 3-9//. usually 4-3 fi in diameter with frequent irregular mycelial 
bodies; Sporangia terminal or intercalary. 7 25 ^ diameter, germinating by germ 
tube; Oogonia smooth, terminal rarely intercalary, spherical 18-26 ^ average 
22\3^± 0*09 borne on laterals 1 5- 90 /a long. Antheridia one per oogonium, 
monociinnus mid arising from immediately below (be oogonium (hypogyiious) 
(fig, 2C"-j jL Anthcridium tube- like, swollen, curved with apex tapering towards 
oogonial wall 5-8 jj. x 11-23/2. Ooospores aplerotic, single spherical 12-23 average 
18*3 /a i 0*00 in diameter, thick walled. Isolated from Snlamtm tuberosum. 

.Middletou (1943). in concurring wilh Drcchsler (1927) for the maintenance 
of Pythtum ulthntiui as a distinct species, points out that its typically monoclinous 
swollen antheridia, which originate close to the oogonium, and curve sharply 
upwards lo make a narrow apical contacr with the basal portion of the oogonium, 
distinguish it from /'y/A//ym rfelnn-yanuiu The^e features were eharaeteristic Of 
the isolate described and figured here. However, in anotl)er isolate, one from 
peas (Isolate !) at hast two types of autheridial structure occurred in the one 
culture. One or more diclinous, ion^ stacked, crook -necked antheridia typical of 
Pylhiuin- dcharvflnuiH occurred in the same culture as monoclinous, suitress 
hypogynous antheridia characteristic of Pythium ulthnum. There were also 
many gradations (fig. 2). 

Middleton, while he refers to the infrequent occurrence of diclinous 
antheridia in Pythium ulthnum says that "they are not distinctive enough to be 
utilized in the. identification of the species." On the other hand. Van Luyk 
(1934), with others, doubts whether such differences between the autheridial 
characters as are m pinioned above are suilicient to segregate the two species, lie 
considers that Pylfiitmt debaryauum. among its different strains, exhibits differ- 
ences of the type described; a view with which the writer concurs. 


Pythium polyvtotphon Sultris 

Hyphae 5-7-jia in diameter. Sporangia spherical, subsjiherical terminal or 
intercalary 7-33/* in diameter and germinating by germ uirm, Oogonia spherical, 
terminal or intercalary, smooth but sometimes with one or more papillae (fig. 3, 
J. M. ) U-25/a av. 16"2/x.±0'22 in diameter. Antheridia 1-2 per oogonium mono- 
dinons, antheridial stalk typically falcate or sigmoid or with curved antheridial 
cell (fig. 3. K.L. M). Antheridial cell not greatly aVVOlfeK and with a long fertili- 
zation tube. 8 r 20/i long x 5-7//, wide. After fertilization lhc apical portion is 
rlivided from the base by a narrow ring at junction with colonial wall (fig. 3, 
F. G. IT,). Swollen diclinous antheridia also occur but mostly in association with 
the smaller oogonia (fig. % J. E. O.). Oospores aplerotic, 7-Z6 p$ av. 13*4 ±0-27 
in diameter with somewhat thickened wall. Isolated from Pk'tim sativum. 

The characteristic feature of this fungus is the shape of the anthcridud 
branch, which may be curved like the Tetter C or double curved like ihe letter i>. 
Ju out* isolate the origin of the antheridial stalk was always monotonous, arising 
some distance from the oogonium and generally curving in ;a wide how to meet 
the oogonial wail in ihe way illustrated by Sidcris. 

The swollen diclinous antheridia applying to the smaller oogonia described 
above suggested the possible presence of more than one si rain in our culture. 
However, it was possible to secure growth from a single hyphal tip of this isolate 
and the cuhnre derived behaved in the way already described. 

The isolate we describe in slightly different from P. polymorpltun described 
by Middleton and appears io be the first record of the association of this fungus 
with risitrn sativum. Previous records associate P. polymorphon with Ananas 
comosus in Hawaii and Nicotiatia Tubaatm in the LJni led' States. 

Pythium vcxattx Do 7'iary 

Jiyplme 2-7 ft in diameter with irregular mycelial bodies associated (Jig. 4. 
JVj, Sporangia terminal or intercalary, spherical to oval 10-26 jk in diameter" and 
germinating by germ tubes < fig: 4, M). Oogonia spherical terminal, 15-26^ av. 
21*7/t±0'10 in diameter. Antheridia 1-2 per oogonium, diclinous. Antheridial 
cell very variable in shape (fig. 4 f G. N.J.O.). In some cases, cylindrical and 
hardly curved or swollen, in others slightly swollen or 50 swollen that die shape 
ii nearly spherical. The antheridial cell or its stalk may be so curved that applica- 
tion to ihe oogonial wall is narrow. In profile it may' appear bell-shaped, and in 
another plane as an allanloid or disc-like structure covering- the oogonium (tig. 4, 
G- 0-). The cylindrical antheridia are 11-18 ^ long by 7-11 y^ wide, the spherical 
7-9 }k in diameter. Oospo res smooth, aplerotie, 1 3-24 //. a v. 1 S ■ 8 n ^ ■ 1 1 i u 
diameter and with a thick brown wall. Isolated from Lycoperxicon vscuicntum. 

Middleton states that "Pythium vc.vmts is readily distinguished from m 
congeners possessing aplerotir. oospores by its typically inonodinous, stalked 
autheridium which arises in close proximity to the oogonium and by the clavate 
antheridial cell with the apex hell-shaped and broadly applied, sometimes fused 
with the oogonial cell/' 

in respect to the origin of the antheridia Middleton fay$ that they are rai'elv 
diclinous, but Dreehsler ( 1946 h), who cultivated Pythium complect ens Brauri. 
and like Middleton considers the name a synonym of Pythium, vexans, remarks- 
that "frequently the mycelial connection between the oogonial sialk and antheridia* 
branch is too remote to be traced for certainty amid the confusion of ramifying 
hyphae." Thus the apparently diclinous character of antheridia in our isolate h hot, 
in itself, sufficient to distinguish it from Pythium vexuns.- Antheridial shape in 
our isolate was very variable. It is true that in certain aspects, trumpet- shaped 
sstructnres flaring out at the region of attachment, as described by Braund, bell- 


shaped apices as mentioned by Middleton, and the bilobate and biramous male 
cells, figured by Drechsler, were observed, but the antheridial cells were by no 
means uniformly of these shapes. 

However, there is another feature in which our isolate resembled descrip- 
tions of Pythiuni vi'xans. According to Middleton the germination of oospores 
of Pylhium is not common; it occurs more readily in Pythiuni vexans, a fact also 
noted by Butler (1907) and Drechsier (1946)' The oospores of our isolate 
germinated fairly readily to produce sessile sporangia winch in turn produced a 
germ tube (fig. 4, K), The sporangia, when they germinate first, produce a 
vesicle into which the contents of the sporangia passed. The vesicles theft 
developed a germ lube. 

Gknkuai. Discussion 1 

Middleton considers that the origin and morphology of the aniheridia afford 
valuable criteria for specific identification. Our isolates, because of close 
resemblance in other respects, have had Lo be looked at particularly in respect to 
antheridial characters, but we found these to be rather more variable than Middle- 
ton's descriptions might suggest. In one case (isolate 5) differently shaped 
antheridia were even found on one branch. (See also the discussion under 
Pythium uliimum.) Although we have distinguished our isolates in the terms 
discussed above,, we should bear in mind the advice of Buisman (1927) not to 
define species too strictly: "It is not at all easy to determine if a special isolation 
belongs to the well-known Pylhium dcbaryanuvi or not/' It is clear, too, thai in 
view of the methods of reproduction concerned and the obvious chances of 
hybridization, intermediate forms might be expected. 

In view of the foregoing remarks, the following scheme of relationship of 
the isolates examined is suggested, remarks refer to salient antheridial features: — 


Pxthium nitvjntni Pvth'mm polynwrphou 
(Potato 1) 

Moriocluious Monoclinous curved 

swollen, stalkless seldom diclinous 


both debaryanum and 
ukimum types 

Pea (6) diclinous 

debaryanum type 

Potato (2) 

Pyt hilt m -vexans 

very variable 

Pea (5) diclinous 

Pea (2) Pph (3) diclinous 

diclinous debaryanum diclinous debaryanum debarvaoum type 
type type 


^ The morphology of nine isolates of Pythium spp. grown in pure culture on 
various media have been studied. Four isolates from infected pea seedlings and 
one from a potato affected by "leak*' disease were identified as Pylhium 
debaryanum Hesse. A fifth isolate, from peas was identified as PvUmnn pnly- 
marphan Sideris. A second isolate from potato with "leak" was identified as 
Pythium ultimum Trow and an isolate from a diseased tomato seedling as 
Pythium vexans De Bary. 

All these species have a number of characters in common; spheroidal 
sporangia and smooth aplerotic oospores, and arc considered to be closely related 
to one another, and this is discussed. 


I wish to express my thanks to Professor J, A. Prescott, Director of the 
Waite Agricultural Research Institute, for affording me facilities and the hospi- 
tality of the. Institute during a three months' visit as a guest. J am also very- 
grateful to Mr. D. B. Adam of the same Institute, who suggested the problem 
and gave a great deal of assistance in preparing this manuscript. Thanks are 
also due to other members of the Institute staff, especially those in the Depart- 
ment of Plant Pathology who helped in many ways. 


Eraun, H. 1924 "Geranium Stemrot caused by Pythium camplcctens n. sp." 
Journ, Agric. Res. ? 29, 399-419 

RnAUN, II. 1925 Comparative Studies of P. de Baryamnn and two related 
Species from Geranium." Journ. Agric. Res., 30, 1,043-1.062 

BursMx\x, C. J. 1927 "Root Rots caused by Phycornycctes." Eaarn. 

Butler, E. T- 1913 "Pythium dr Baryamtm Hcssc." Mem. Dept. Agr., India. 
Bot* Ser. 5, 262-266 

Butler, E. J. 1907 "An Account of the Genus Pythuun and some Chytri- 
J diaceac." Mem. Dept. Agric. India, Bot, Ser. 1. 1-160 

Drkchsler, Ch. 1946 "Pylhium vJtimum and Pythium debaryanum. (Ab- 
stract.) Phytopathology, 17, 781-864 

Drechsler, Ch. 1946 "Several Species of Pythium peculiar in their Sexual 
Development." Phytopathology, 36 v 781-864 

Johann, H. 1928 "Grated Carrot Agar favourable for Studies of Pythium.'' 
'(Abstract.) Phytopathology, IS, 710 

Li:yk. A, van 1934 ''Pythium de Baryovum Hesse emend, de Bary." Beded. 
Phytopathol. Lab. W. C. Scholten. Baarn, 13, 23-2S 

Matthews, V. D. 1.931 "Studies on the Genus Pythium/' Univ. of N. Caro- 
lina Press. 
Middleton, J. T. 1943 "Tbe Taxonomy, Host Range and Geographic Distri- 
bution of the Genus Pythium." Mem. Torrey Bot. Club, 20, 1-171 
Scuulz II V 1942 "Arbeitsmethoden bei Kultur- und Infektionsversuchen 

'mith Pythium Artcn." Centralblatt L Bakt, II, Abt. 105, 248-254 
Sideris, C. P. 1932 "Taxonomic Studies in the Family Pythiaceae, II Py- 
' thium.' Mycologist, 24, 14-61 

Trow, A. II. 1901 "Observations on the Biology and Cytology of Pythium 
ultknum n. sp." Ann. Bot., 15, 269-312 


ByD. Mawson 


The sediments now to be considered constitute a very thick series of formations of late Precambrian 
age. Though varied somewhat in nature and in mass from their counterparts in the type area, they 
are the northward continuation of Howchin's Adelaide Series of the vicinity of Adelaide. 
Deposition took place in a great geosynclinal trough. 



By D. Mawson * 
[Read 9 October 19471 


t. The Gkosynclin'ai. Hasix of Accumulation , 260 


A. General Remarks .. ., «> .. . ~,« 

B. Proterozoic sediments exposed to the west of Copley . . 26? 
1. Tabulated sequence of strata from the glacial horizon to that of 

the Brighton Limestone . . / -. 

1. J. he strat^raplncal equivalence of the Mt. Arooiia Quartzite 

with the Aidgate Sandstone .. , . 2fA 

C. Proterozoic sediments of the neighbourhood of Quurn .. 265 
J. General Remarks 

2. Section from Devils Peak 1o Lake Torrens StmWiaiul .. " fL 

3. 'Hie Proterozoic sequence in the neighbourhood oi" Mundallio 
Creek . . . . , , 

(a) The Emeroo Range basal rudaceous and arenaceous forma- 
tion .... 

/I \ TL , 1 "* ** M • * ^ /U 

(b) 1 he ao-omiie-magiiesite series 2"0 

(c) The glacigetie sediments .. .. " ?71 

(d) Laminated (Tapley Hi'l) postglacial argillites .. .. ~ T i2 

(e) The Brighton Limestone horizon " ^ 

(1) Argillaceous and arenaceous sediments above the Brighton 

Limestone horizon , . . p . t ._ 

(g) The Pound Quartzite (Cambrian) ' 27> 

D. Bnef review of the several major formations constituting- the Pro- 
terozoic record of the Copley and Quorn Areas and the Pound 
Sandstone *■-..... ^, 

E. Outcrops of the basal quarlzfite elsewhere in the South Flinders 

Causes „ 

1. The great quartzite of the Port Gcrmein Gorge .. .. 2 7> 

2. The massive quartzite of Mt. Remarkable 2 *~4 

F- The Mt. Aroona-Emeroo Range Quartzite as distinct from the Pound 

Quartzite , 

G. Transgrcssional Proterozoic sediments located to the west of the Lake 

Torrens Sunkland „. 

1. The Tent Hill Formation 275 

(a) The Corunna Range rudaceous and arenaceous formation .. 275 
(h) Equivalence of the Corunna Ratine formation with that of 

the Emeroo Range , . , , . 275 


IV. Other Correlations ..... v,^ 

• - - ■ . . 27/ 

V. Summary „ 4 , 

VI. ., .. „~ 

•• •- .. 278 

The sedimetiLs now to be considered constitute a very thick series of fov„ 
tions of late Precambmn age. Though varied somewhat in nature and m mass 

on a- 

* Geology Department, tlhivfrgftjt of Adelard 
Trans. Ko>. Sec. S. Aust., 71, (2), 1 December, 1947 


frron their counterparts in flic type area, they are the northward continuaiHw 
oi Tlowchin's Adelaide Series of the vicinity of Adelaide. Deposition look 
place in a great gcosynclina! trough. 

The geological record evidenced in the Mount Lofty and Flinders Ranges 
ck-a.Iv demonstrates the existence in later Precambrian and early Cambrian 
\jmf% pi a geosvnclinal depression extending from south to north for a d.stance 
of several hundred miks, then turning' to north-west through the region i if the 
Wiilouran Range on to the Auiadeus Strait region via the Mount Denuu- Margaret Range and the Everard Ranges. In Central Australia it appears 
to have been mainly located between the great belt of basal conglomerate repre- 
sented by Avers Rock and Mount Olga on the south and the MacDonnell Ranges 
on the north. From there it may have extended in the north-west tQ the region 
of the "Desert Basin" and the Kimberley Ranges. 

On the western margin of this great orogenic feature was an ancient massif 
constituting the Early Precambrian land area which has reccved the palaeo- 
ceo K raphk name of Yilgarnia (Cotton 1930). From the denudation of Y.Jgarn.a 
There came much, probably most, of the dctntal material contributing to ti.e sedi- 
ments of that, progressively subsiding geosyuclinal trough. 

The eastern margin of the basin is not so clearly defined, for there is still 
some uncertainty as co the age of the rock* of ihe more im'tamorpluwed ten am 
us the eastern hank* of the Mount lofty Ranges Further north, where the 
scdimsrdi of the geosynclioal depression are easily recognisable, I hey tor the 
mofet part, pass beneath the Tertiary formations of the Cult and 
tbc Mesozoic succession occupying the basin of the one-time Lake VVallu..... 
(July in the Barrier Ranges of western New South Wales and in the neighbouring 
reoiml of north-eastern South Australia and possibly also the nortli-east tip ot 
i he Hinder* Ranges can we clearly demonstrate the existence of Precambrian 
land on the eastern side of this tffte Precambrian depression. This is Thy at. S 
('1932) "Eastern Massif" and Andrews' (1937) 'AVillyamia." 

In the northern Flinders Ranges, the swing of the bedded sediments around 
to the west and north-west appears to have been due to the existence of a more 
stable crustal block to the north and north-east. This is detm.tely so still fartttgr 
•north, in the region of the MacDonnell Ranges. 


Returning now to the region which we have studied in considerable detail 
.bat hKween 'the ancient land masses Yilgarnia and Witlyarma. During the past 
40 vears I have had many opportunities of examining these ancient geosynclma 
sediments in selected areas in the region occupied by the Flinders Ranges and 
eastward to the Barrier Ranges. Measurements have disclosed that in the centre 
of the basin the total thickness of such deposits is immense (Mawson 1!»Z}. 
In the marginal regions, some at least of the very numerous formations repre- 
sented thin out or are absent. Already much relating to the thickness and dis- 
tribution of the sediments of the deeper parts of the trough has appeared ma 
series of contributions submitted by mc to this Society, but there still remain* 
n be published extended observations on the glacial and lluvioglacia succe^imi 
,,! the central portion of the basin. Thus far. 1 have dealt .with the types ol 
raiments represented and their respective thickness. The.r petrographic 


characters have been only very cursorily touched upon, a detailed account of such 
having been purposely withheld until the final review when the varying phases 
of sedimentation can be comprehensively dealt with. 

Concerning the corresponding 
sediments represented in the eastern 
region bordering the Willyamia 
massif, I have as yet published 
(mainly D. M. 1912) only very- 
limited detail, reserving most of 
the results of work in that region 
for consideration in relation to the 
sedimentation of the geosynclinal 
basiu as a whole. In that region 
the extensive survey conducted 
under Andrews (1922) has added 
greatly to the knowledge of that 
area, though principally relating to 
the Willyama series itself. 

Notable contributions relating to 
areas of (he central and western 
belts have been furnished by IIow- 
chin and by members of the South 
Australian Mines Department, in- 
cluding, more especially. Dickinson 
(1942), Jack (1914, 1922), and 
Segnit (1929 and 1939). 

Further south, in the Adelaide 
region, Howchin has in the past 
been the chief investigator of the 
stratigraphical succession, but more 
recently Madigan (1927), Hossfeld 
( 1935) , Barnes and Kleeman 
(1934), Segnit (1937) and Sprigg 
have published important contribu- 
tions to this subject. The latter, 
who as a student, assisted me on 
occasions in field Avork conducted 
hi (he Flinders Ranges, undertook 
and has now completed a check of 
Howchin's original work in the 
vicinity of Adelaide. He has pub- 
lished (1942 and 1946) an account 
of the various formations consti- 
tuting Howchin's Adelaide Series 
as occurring in the type locality, 
including thickness and general 
distribution in that area. 

Locality Plan 


When examining an important occurrence of magnesitc near Copley which 
is located towards the western side of the great gcosynclinal trough, 1 was struck 
by the regularity of the sedimentary formations and freedom from faulting. 
Accordingly, a traverse of some 16.0*00 feet of these beds was made extending 
upwards from an immensely thick arenaceous formation to somewhat above 
a glacial horizon, correctly regarded as equivalent to the Sturtian Precambnan 
glacial horizon near Adelaide. This belt of sediments, which dips steeply to the 
east, was fouud to extend for miles along* the .strike without a break. From 
Mount Aroona it was traced for a few miles to the south-east, crossing to the 
east of the main north road to Copley: to the north-west it extends for more than 
20 miles from Mount Aroona, passing' through Myrtle Springs Station and on to 
the neighbourhood of Termination Hill. There major faulting and shattering 
abruptly dislocates the formation. 

U FPCn ADILAOE 5v«IEE hear cqt-le-y 


- — I ***Wtfjj 

Fig. 2 

A detailed section of the lower members of the succession has already been 
published (Mawson 1943). A further geological section is now suhmitU'd as 
fig! 2 herewith. This carries the sequence from the glacial stage to the Brighton 
Limestone horizon. The cross-section illustrated refers to observations made in 
a location situated some two miles northward along the strike, from that of the 
earlier Lraverse. In the locality indicated beds higher in the sequence than 
exposed on the former line of section are available for study. 

It .should be mentioned that the suggested hidden fault at the eastern end 
of the earlier published section almost certainly does no: exist. There die out- 
cropping Precambrian rocks cease and the low-lying open country beyond is 
occupied by the Leigh Creek Triassie basin. According to earlier reports rdatini; 
ta this coalfield area the Triassie strata occupy a trough-faulted belt- Recently, 
however, a survey of the area made by officers of the S.A. Mines Department 
Ufr* shown that the Mcsozoic rocks occupy basin-shaoed. but not faulted, depres- 
sions in the older forruaiions. 


Tabulated Sequence, as displayed in Fig, 2, of Strata from the 
Glacial Horizon to that of the Brighton Limestone 

Pre-Crlacial Sediments 

1 32 ft. of Dolomite: strike N.20 a W. (true), dip So° easterly. 

2 158 It. mainly shales. 

(a) Flaggy calcareous shales Thickness 104 ft. 

(b) Yellow dolomitic shale, strike N. 30° W, dip 60° easterly. Thickness 
33 ft. 

(c) A sandstone band. Thickness 15 inches. 

(d) Sandy shale. Thickness 20 ft. 

Glacial and Fluvio glacial Sediments 
(cf. items 32 to 36 of the 1941 section.) 

3 116 ft. of Fluvio-glacial muds and siltstone. 

(a) Sandy fluvio-glacial grit with erratics up to 5 inches diameter, 
Thickness 10 ft 

(b) Argillaceous siltstonc. Thickness 48 ft. 

(c) Fluvio-glacial mud (with gravel bands). Thickness 13 ft. 

(d) Fluvio-glacial mud of coarser texture and with occasional erratics. 
Thickness 8 ft. 

(c) Laminated silt with a little grit: strike N.35°W, dip 64° easterly. 
Thickness 11 ft. 

(f) Fluvio-glacial muds. Thickness 26 ft. 

4 299 ft of Tillitc. 

5 143 ft. of Laminated Fluvio-glacial series. 

(a) Laminated shale with occasional erratics: strike N.42°W, dip 46" 

easterly. Thickness 80 ft. 
(h) Laminated shale with erratics up to 6 inches diameter. Thickness 5 ft. 

(c) Finely laminated fluvio-glacial mud and thin hands of grit. Thickness 
58 ft. 

Post-Glacial Sediments 
tf 2,016 ft, of laminated flaggy shales, etc, 

(a) Laminated shales. Thickness 110 ft. 

(b) Laminated shale with some calcareous hands. Thickness 30 ft 
(c.) "Laminated shales. Thickness 106 ft. 

(d) Flaggy shales; N.35 G W t dip 4S° to the east Thickness 567 ft. 

(e) FaintU laminated fissile shales; dip 40° to the cast. Thickness 307 ft. 
(!) Slaty mudstone. Thickness S3 ft 

(g) Flaggy slate; strike N.35°W ( average dip 3S a to the east. Thickness 
843 ft. 

7 724 ft. of somewhat calcareous flaggy slates. 

(a) Flaggy calcareous slate in beds a few feet in thickness inlerbedded with 

slates. Total thickness 43 ft. 
(lb) Flaggy, faintly laminated, slightly calcareous slates; dip 29° to the east, 

Thickness 681 ft 

8 14 ft. of flaggy limestone. 

9 248 ft. of slates. 

(a) Laminated flaggy slate. Thickness 100 ft. 

(b) Thin-bedded flaggy slate; dip 15° to cast. Thickness 148 ft 

10 2i ft. of Impure Limestone. 

11 94 ft. of slate and sandy argillaceous Hags. 

(a) Slate; dip 15° to the east. Thickness fi2 ft, 

(b) Sandy argillaceous flags with some calcareous bands. Thickness 32 ft 

12 500 ft of flaggy calcareous slates; dip falling off from 10° to 5° toward the east. 

13 Massive limestone dipping at a very low angle. This is the Brighton 
Limestone horizon. 


From item (3) to item (12) inclusive, representing all sediments from the 
base of the Sturfiun glacial formation tu the base of the Brighton Limestone 
amounts to 4,176 feet 

Toe STitArtoKAPHTc.M. Equivalence of the Mou.vr Aruojna QtjArtzitk 
with the. Aldgatk Sandstone 

The succession of strata as exposed to the west of Copley (Mawson 1941 ) 
cxhibils sufficient correspondency with that below the Sturtian Tillite horizon in 
the neighbourhood of Adelaide, as established by Ilowchin, to indicate equiva- 
lence in ti_me of these respective series of beds. In the neighbourhood of 
Adelaide, below the Sturtian Tillite, in descending order arc laminated shales 
and minor quartzites, then shales with many intercalations of dolomite (Beaumont 
Dolomites) more argillitcs. some phyllitic in places, and then the Mount Lofty ov 
"Thick" quartzite. 

As. this is broadly the succession at Copley the Mount Aroona quaruite 
was accepted (Mawson 1941) as the equivalent of HmvduVs Thick Quartzite. 
This decision was based on the fact that at Copley there is no outstandingly thiek 
quartzite in the series other than that of Mount Aroona; also at that time there 
was a grave doubt as to the. accuracy of Howchin's succession below his "Thick 

It will be recalled that Jlowchin. on account of faulting and structural com- 
plications, was not able, in the region between Mount Lofty and Aldgatc, to 
satisfactorily relate the outcrops of the lower members of his succession, lie 
finally relied for sequence in that part of his series upon the succession which 
he established in the Torrens Gorge area, which is a region greatly affected by 
faulting and structural complications. 

Later work (Barnes and TCteeman 1934 and Mawson 1939 and 1946) had 
cast some doubt upon the relations, as established by Howcbm, of the Torrens 
Gorge succession below the Thick (Juartzite. The suggestion made was that 
Howchin's beds below the Thick Quartzite, as interpreted from the Torrens 
Gorge outcrops, might represent a duplicarkm by faulting of his Upper Phyllites 
and Dolomites. The probability that such is the case was favoured at the time 
of publication of the Copley section (D. M. 1941). 

Smce then, Sprigg (1946) has completed his reconnaissance of the Adelaide 
Series in the neighbourhood of Adelaide, and though not entirely satisfied with the 
evidence, owing to faulting and displacements in the Torrens Gorge area, he 
concludes that the Upper and JLower Torrens Dolomite formations (Howe-bin'* 
Upper and Lower Torrens Limestones) sequentially follow above a basal ilmemtic 
sandstone and below the Thick Quartzite. 

Howchin recorded that his Upper Torrens ''Limestone" (actually dolomite) 
had associated with it bands and nodules of chert. Now Sprigg has foimd magnetite 
associated with this Upper Torrens Dolomite. As both these chert and magnesite, 
are peculiarities of the mam dolomite series near Copley, it is now evident that the 
dolormte-magnesite series above the Mount Aroona quartzite should be correlated 
with the Torrens Gorge Dolomites, which latter are now accepted (Sprigg 1946) 
with confidence as stratigraphically below the Thick Quartzite. Thus it is that 
Mr. Sprigg (1946, p. 328) has indicated that the Mount Aroona Quartzite "may 
provti to be the equivalent of the basal ilmenitic sandstone in Howchin's type 
area, while Howchin's Thick Quartzite would probably be contemporaneous with 
one or more minor quartzites in Mawson' s Magnesite serves of the Flinders 

I agree with this suggestion, a conclusion eou.seque.tu on Sprigs proof that 
the Torrcns Gorge dolomites are a separate and earlier magucshiferous series 
distinct from the Beaumont Dolomites (Howchin's Bine Metal Limestone). 
Evidently, the Mount Aroona quartzite corresponds to the Aidguie sandstone; 
that is. with the basal member of Howchin's Adelaide Scries. Howehin regarded 
the Aldgate ilrncnitic sandstone as of about 200 feet in thickness. Spring finds 
that, in {he Aldgate area the first 100 feet of lhe Adelaide Series is ilrncnitic sand- 
stone, and that while the relations are not quite clear this appears to be followed 
by upwards of 2.000 feet nf alternating sandstones and gritty and sandy slates, 
all of wlu'ch are located, below the horizon of the Lower Torrens Dolomite, 


G^mkr'w. Remarks 

Subsequent Lo my examination of the Copley area, <m important occurrence 
of magnetite was reported in the neighbourhood oF Mundallio Creek, west of 
Quorn, situated about 130 miles in a nearly due smith direction from the former 
locality (see fig. I). This has proved to be a repetition of the formation at 
Copley, though containing less dolomite and magnesite. Here the general 
sequence of formations is the same as at Copley, hut here the upward range 
exposed is greater. An unbroken succession extends from a great basal rudaceous 
and arenaceous formation, equivalent to the Mount Aroona quartette of the 
Copley ^ area, up to and through the basal sandstone (Pound Sandstone) of the 
Cambrian. There is thus exposed a complete succession corresponding to the 
Adelaide Series. 

Howehin (1928) recorded some ot the geological features of the neighbour- 
hood uf Quorn with special reference to the tillite, but he did uot recognise the 
magnesite formation, nor did he locate the remarkable development of quartzite 
and conglomerate which constitutes the basal member of the Proterozoic sequence 
in (hat area. More recently, Scgnit (1939) completed a geological map of some 
M square miles of that area, but missed recording the existence of magnesite. and 
mistook the enormously thick basal quartzite of the Proterozoic sequence for 
(he Pound Sandstone-Quartzite which is some 13.000 feet stratigraphicaJly 
above it. 

3f^ I \ __ 

Fig. 3 

Here again, as in the case of his map of the Mount Scott - Mount Aroona 
area near Copley, Segnit (1939) mistook the thick basal quartzite of the Pro- 
terozoic for that far above in the sequence ushering in Lower-Cambrian 
tossiliferous formations. J lis interpretation necessitated the incorporation in his 
map of a grand strike-fault along the face of the quartzite on its eastern side. 
We found no evidence of stielt a major fault, though small displacements do occur 
and would be expected at the contact of so great a mass of quartzite with the 
less competent overlying argillaceous beds, where orogenic forces on a grand 
scale have thrown the sedimentary formations into a series ni folds. 


Elsewhere in his map, Mr. Scgnit unconsciously introduces faults to recon- 
cile, it seems to me, the observed outcrops with a preconceived conception of the 
sequence. Unquestionably, there are a number of large scale faults in that area. 
A notable strike fault passes through the quartzite belt of the Devil's Peak as 
mentioned by Howchin (1928) and illustrated by Segnit {1939) ; another trends 
along the valley between the Devil's Peak and the Dutchman's Stern Range; 
another throws "down the country to the west of the great basal quaitzUe. shew- 
ing it in places, Other apparent lines of faulting are indicated in the diagram 
herewith. Minor faults and displacements are, of course, numerous, as would 
be expected~in a region at one lime (late Cambrian ?) sub jeered to an alpine 
orogeny and subsequently, after peneplanation, recreated once again as a range 
of mountains (late Tertiary) of the nature of block uplifts. 

Suction from Devti/s. Pkak westward to the Lark Torrens. Suxkland 

The general relation of the strata in this area is illustrated in fig. 3 herewith, 
which is a section from the Port Augusta- Lake Torrens Sunkland across the 
Ranges to the Devil's Peak, We have found that the field distribution of out- 
crops of the various formations is fundamentally dependent on the development 
of a system of pitching folds. The structures thus developed are modified by 
some major and frequent minor faults. 

A typical anticlinal fold is that to he observed at Pichi-R:chi Pass in the 
neighbourhood of the Devil's Peak. There the great Cambrian quartzite (Pound 
Formation) of the Dutchman's Stt-rn Range descends again to the east as the 
DeVils Peak. The intervening valley region of Pichi - Richi Pass is occupied 
by sequentially underlying sub-Cambrian beds which, however, arc sliced by a 
strike-fault along the crown of the anticline with considerable downthrow on 
the east side of this fault. Another strike-fault on a considerable scale slices the 
quartzite of the Devil's Peak mass as illustrated in fig. .1 The quartzite mass 
of the Devil's Peak is truncated to the north by a fault cutting diagonally across 
the strike, while to the south it can be traced looping around on a course some- 
what checkered by faulting, but eventually joins up with the southern extension 
of the somewhat attenuated Dutchman's Stern Uange. What is important is that 
outcrops ot the Pound Quartzite (Segnit'a D7 formationj shown on Segmt's 
map a* appearing in the Pichi -Richi Pass region between its outcrop in the 
Dutchman's Stern Range and its repeat in Devil's Peak arc not portions of the 
Pound Quartzite proper but are underlying greywackes, arkoses and shales 
separated sequentially from the former by a thick formation of chocolate shale. 

In this region the sequence of beds from the hasal conglomerate ot the 
Proterozoic formation to the basal Cambrian quartzite is well illustrated in a 
Straight run of beds on the western side of the Ranges. There is there a 
minimum of disturbance by faulting, a feature strikingly indicated in Segmt's 
(19.39) coloured plan of the locality. As already stated, however, we do not 
agree with him where he introduces a strike-fault on a grand scale along the 
upper limit of the hasai ijuartzite. The latter wc have found 10 be equivalent ot 
the quartzite of Mount Aroorui in iKe Copley area. 

TbiK Pkoterozotc Succession tn the Neigh uourfjgod tir 
McNTMruro Crf.ek 

The succession of beds in this locality is graphically represented in fig. 4 
herewith. Descriptive details of the individual formations are given bdow. 
Neither the absolute bottom of the great basal coHKlomerate-quartxite formation 
nor the nature of the pre-existing; (underlying) rocks is demonstrable in this 
area owing to major block faulting of Tertiary times responsible for downthrow 
of 6 wide belt of country (Port Augusta Lake Torrens sunkland) lying to the 
west of the Emeroo Range. This problem is discussed later. 



Tabular Statkment of the Pkotukgzoic Succession at Mundallio Ckeek 

The Proterozoic succession of strata occurring in the neighbourhood of 
Mundallio Creek, St}uthern Flinders Ranges, The beds are numbered in upward 
sequence from the lowest exposed beds. True bearings given in all cases. The 
sequence is graphically displayed as fig. 4. 


1 Basal beds of course water-laid conglomerate occurring in an alternating- 
succession with finer arenaceous sediments. Pebbles met with tfp to 6" 
in diameter, mainly constituted of liver-coloured quartzitc, but some 
appear to be (Jawfer Range porphyry. Dip 55° E, strike N.G° K. Near 
the upper limit of this section is a sandstone with marked ilmenjtic 
banding but free from pebbles. Strike N. 4 Q W, dip 55° E. The base of 

this section was not observed but the total thickness measured is - - 600 

2 Bands of iirnenite-bearing quartzitc- alternating with conglomerate, each 
component band being about 3 ft. thick. Strike N. 4° W, and dip 70° E. 62 

3 Alternations of sandstone with reddish argillaceous partings each of the 

latter ranging from 2 to 3 feet thick. Dip 70°. - 270 

4 Quartzite without any shale partings and with only faint traces of 
bedding laminations. All but near the base is notably white and hard. The 
dip ranges from just about 70° below to near w at the top limit. The 

strike swings around locally at the upper limit to N. 4° W. - 490 

5 A belt of sandstone somewhat reddened in part and with traces o[ minor 

shearing. ------------- % 

6 Quartzitc of a generally uniform character with dip to the east between 

72° and 65°, Strike N.14°W, below to N\ true in the upper [beds, 1,276 

7 Quartzitc, hard below and less resistant above. Average dip 70° - 1,294 
The Emeroo Range basal rudaceous and arenaceous sediments: - - Total, 4,288 

8 Soft sandstone in part argillaceous, Outcrops deficient. Dip 72° to the 

eastward ------------- 262 

9 (a) Sandy shale, Dip 70°. - 144 

(b) Grey dolomitic mudstune -------- R7 

(c) Flaggy sandstone and marly micaceous shale ----- 42 

(d) Sandy Marl ..„-.--- 87 

(e) Grey marly shale. Strike N. true, Dip 73° E. - 70 430 
S*ndy to marly transitional beds: -------- Total, 692 

10 (a) Two very narrow scams of nodular magneske embedded in marl 11 
lb) Poor outcrop, but apparently marl and marly shale with some 

narrow seams of magnesite. --------42 

(c) Shales below with thin dolomitic bands above: then sandy marl to 
marly shale with thin dolomitic bands above. ----- 54 

(d) Hard dolomite. _______-_- .8 

(e) Marly Shale. ----------- 39 

ff) Bands of dolomite and thin-bedded marly shales. - 219 373 

11 (a) Marly shales with several magnesite Bands. Dip 70° 17 
(b) Banded dolomite with irregular seams of chert and magnesite 

passing above to alternating beds of shale and dolomite. Magnesite 
mine workings are located in this section. ----- 329 346 
Marly series with dolomite and magnesite: ------ Total, 719 

12 Arenaceous beds enter the succession, becoming dominant over the 
dolomitic shales at the top of this section where there appears a band of 
flinty quartzitc exhibiting contorted bedding ------ 

13 (a) Laminated sandstone becoming more massive above. , - - 139 
(b) Carbonate laminations appear in the sandstone and it finally passes 



into a calcareous sandstone (carbonate cement). - 122 

(c)'No outcrops: probably soft calcareous sandstone. - 115 376 

14 (a) Sandstone which in part has a dolomite cement and occasional frag- 

ments of dolomite embedded in it, ------- 82 

(b) An association of marly sandstone and sandy marl, strike St* true 

dip 63° E. - ------- 212 

(c) Marl ------------- 01 

(d) Laminated shale. ----------78 

(ej Hard uuartzite. strike \\4 D W. dip 65° E. - (54 497 

15 (a) Marly shale. ----------- 67 

(b) Sandy dolomite. ---------- 

(c) Sandy shale with occasional narrow scams of dolomite. - - 71 

(d) Strongly laminated (varve-like) shales. _----- 40 

(e) Sandstone with some marly base. Certain beds are mote marly- 40 233 
Sandstone and shale, much of it dolomitic and marly; - Total, 1,392 

16 Laminated shales. .---------- 3i>2 

Laminated Shale: - - - TotaI - 362 

17 (a) True tillite (quite imsortcd ') containing erratics, also fragments of 

chert and dolomite of the underlying series. With tbc tillite are 
some intercalated bands of morainic mud. ----- 62 

(b) Finer grained, nuvio-glaetat beds more firmly cemented and resistant 
in the upper section. --------- 

(c) Irregular sand intercalation? (each a few yards across) ramifying 
through the more regularly distributed fluvto-grlacuil sediments. 13 

(d) Sandy tillite passing upwards into typical lillite for a few yards in 

thickness then reverting- to fluvio-glacial sediments. - - - 25 300 

18 (a) Fhrvio-gtacial, well laminated slates. ------ 539 

(b) Slate (rock- flour type) poorly laminated. Strike N. true, dip 6(1° E. 167 706 

Series of Glacial and Glacigene sediments: - Total, 1,006 

19 (a) Somewhat softer slates. --------- 557 

(b) Laminated (Taplev Hill type) slates; somewhat flinty. Strike 

N,6 n F.., dip 65° --------- 101 65B 

Laminated slates - - - Total, 658 

20 (a) Slates somewhat calcareous, followed by a belt 54 ft. thick of 

argillaceous limestone in which are reddish bands and pellet mark- 
ings not unlike the "hieroglyphic' structure recorded elsewhere in 
post-glacial calcareous strata of the S. Austr. Protcrozoic record. % 

(b) Strongly calcareous edgewise pellet conglomerate. . . <) 

(c) Slates. Strike N. 6° E. dip 75 s E. - - K$ 264 
Calcareous slates and argillaceous limestones; * Total, 264 

21 Slates which weather reddish on outcrop though grey at a depth. Stains 
of copper carbonate obrerved in bedding planes at several points alone: 
the outcrop. This section becomes more arenaceous in its topmost limits 

and there weather reddish brown. - - - 677 

22 Shales and sandy shales with intenbedded sandstones (brownish outt:rv>i>). 

Strike N.6°E. Dip 75° E. - - - w 

Slates with limited arenaceous intercalations: ----- Total, 1,590 

23 Reddish sandstones and some mterbedded shnle formations. - - 1.0 r 7 

24 Soft purple shale. - - - 1 ^ 72 

Purple series of sandstones and shale: ._-__- Total, 2,129 

Total Proterozoic Formation represented, ------ 13,100 

Basal Cambkian Formation" 

25 Reddish to purple sandstone. ----- - 2j\ 

Massive sandstone and quartzite. ------- - 1,83- 

Massive sandstone and quartzite (Pound Formation); - Total, 2,083 


A comparison of the formation;.; as developed in this Mundallio Creek 
section with their equivalents in the Copley section demonstrates the fact that 
in the former area., except in the. case of the basal beds, the geosynclinal Prn- 
terozoic sediments are not nearly so strongly developed as in the latter locality. 
The explanation appears to be that the Mvmdallio Creek area was nearer to the 
nkl Precambrian shore-line, where the subsiding basin was less responsive tu 
isostatic loading. Further eastward, in the Carrietoa - Oraparinna belt, the total 
thfcktttef! of the corresponding series of beds is notably greater even than in the 
region 63 Copley. 

The Emkroo Ranoh Basal Rudaceous a>ji> Arenaceous Formation 

On the line of section across the Emcroo Range (fig. 4) a thickness of 
about 4,288 feet of quartzite and conglomerates arc traversed, of which the lower 
800 fecr ra mainly conglomerate, judging by the coarseness of the conglomerate, 
it is likely that the actual base of the formation is not far below the base of the 

In the case of the Mount Aroona "Range quartzite of the Copley area, we 
found near Myrtle Springs Head Station some small pebble* appearing m the 
quartzite at the lowest exposed outcropping horizon. There, as in theErneroo 
Range, the further downward extension cannot be observed owing to a major 
down-throw of the region to the west. 

The thick rudaceous basal section, so well exposed in the Emeroo Range, is 
composed of grit bands and pebble beds of varying thickness. At some horizons 
bonlders of 6 inches diameter are not uncommon, but generally speaking such 
a size is exceptional. Pebbles of white quartzite and of liver-coloured quartzite 
are commonest, but schist and lumps of ilmcntte arc not uncommon ingredients. 
Some brown pebbles resembled the felsitic base of the Gawler Range porphvry. 
but none such were observed to carry porphyritic feldspar?. 

Towards its upper limit, this basal rudaceous section of the formation passes 
through a stage of alternating beds T each several feci, thick, respectively of con- 
glomerate and of pebble-free ilmeniric sandstone. Then follows a short run of 
similar alternations of ilmenitic quartzites and red siiaie. Thereafter follows 
the main mass of the formation, which is qiiartzite and arkose. 

A penological peculiarity, distinctive of a certain horizon in this quartzite, 
is that of a very obvious banding, due to recurring deposition of clear water- 
sorted quartz grains of larger size than usual along bedding planes. This banded 
quartzite has usually a faint to a somewhat stronger tinge of pink; it also may 
be spotted with white specks of kaolinized feldspar. This latter feature is also 
strongly developed ai some other horizon?., where in many casts the original 
bedding planes are not evident. 

The sand grains of ihe early stages of deposition ot this enormously thick 
formation are generally coarser and more arkosie than in Ihe case of later con- 
tributions. In this section microelme is the most abundant mineral, apart from 
quartz. A pink to brown colouration is usually noticeable in this lower zone. 

The quartzite of the middle region of the formation, where it is apparently 
most resistant to weathering and consequently >kmds out as the highest ridge 
of the range is whiter, more finely grained and not noticeably arkosic. 

The DoLomjte - Magnksite Series 

In the Mnndailio Creek locality this series is developed on a more limited 
scale than in the Copley area, where strata of dolomite and magnesite. (Mawson 


1941) arc scattered at intervals throughout the entire sequence between the thick 
basal arenaceous belt of Mount Aroona and the glacial series above, a total thick- 
ness of sediments of about 7,000 Feci. The outcrop of these beds extends 
unbroken both to the south-east and to the north-west from Mount Aroona for 
a considerable distance. 

Beyuikl Mount Parry, some 5 mile* r.o the north, this formation is still well 
exposed in (he surface. " There rhe strata dip very steeply— nearly vertical— to 
the north-east, Thereabouts the main magnesite-bearing section ot the bed* 
occupies a surface width of 3,000 feet; included (herein are a number of steftnft 
of magnesile, with a total thickness of not less than about 50 feet. The most 
westerly magnesJte outcrop is of the very coarse nodular variety; there is there a 
bed of 7 feet in thickness separated from a second bed 2 feet in thickness by a 
thin parting- Another concentration of beds of magnesite occurs near ihe 
upper limit of the eastern side ot* the formation. 

Details of the original area near Mount Aroona have already been recorded 
(Mawson 1941), but since Lhal date much exploratory mining work by cnstcaiis 
and pits lias been done on a width of 100 yard*, selected where richest in iflPg- 
Tteshe, The result of opening up the outcrop has been to expose more magnesite 
bands than were reckoned to exist when we measured up the original section. A 
very large tonnage of a good grade of magnesite is now exposed. 

T1»e magnesium-rich sediments were obviously laid down in shallow, saline. 
land basins. The magnesih- is uf sedimentary origin as a primary or pene.eon- 
temporaneous sediment. Many of the maguesite beds are. composed of nearly 
pure magnetite, containing extremely little calcium. Occasional beds reach a 
purity trfv9S% MgCO... Wc now know I his magnesium -rich formation to be very 
widespread in South" Australia. It is most strongly developed in the more 
westerly areas, especially discussed ni this paper. Nevertheless, important beds 
also appear in the more central region of the geosynclinal basin, such as the 
neighbourhood of Ralcanoona (where magncsite rock composed of uniform 
pellets was originally described by Fred, Chapman as Mawsonella), Johnburgh 
and Robertstown. Sprigg (1946) has recorded pellet magncsite near Adelaide, 
where ir is associated with the Upper Torrens Dolomites in what is apparently 
this same stratigraphical horizon. 

The most remarkable, feature, of this magncsite is that it usually occurs in 
a peculiar nodular form. A study of this structure has convincingly shown that 
the nodules arc original pellets laid down in beds as in the case of clay pellet 
formations. This appears to have been a development from maguesite sludge, 
which had evidently accumulated m the first instance us thin laminae of almost 
pure magncsite, eventually to he fragmented and re-acaunnlaied as pellet beds. 

Skihmexts ok GuAeiAt. ami Fluvioolactal Orhum 

In sequence above the Dolomite - Magnetic series rorncs a variable thick- 
ness of glacigene beds. These comprise depositions of true tillite, bedded fiuvio- 
glacial mudstones with or without embedded erratics and some intcrglaoal bedded 
arenaceous auci argillaceous sediments, which are on occasions typically varved. 

The variability in thickness of these glacigene beds in Ihe area now \inder 
consideration is considerable. Ju the section near Copley 608 feet of obviously 
glacigene beds were met with, but at Mtmdallio Creek there was fouud a total 
thickness of 1,006 feet. Further sotnh, near Crystal Brook, the section published 
by Howchin (1928) shows, a thickness of 1.240 feet. Further cowards the 
centre ot the basin Ihe thickness increases considerably. However, it is not only 


that the total thickness varies between wide limits within the State of South 
Australia, but what is still more interesting is that the records preserved In 
different localities within the State vary considerably in regard to the number 
of glacial culminations evidenced. In some localities the glacigcrte beds embody 
only one deposit of true tillite. whereas in other places there are 2, and even 3, 
major horizons of tilUte. These are separated by fluvial and fluviogiacial sedi- 
ments, presumably of intcrglacial periods. 

One factor in the irregularity of deposition from place to place is due to 
the fact that, over large areas nf" South Australia, and adjoining territory, the 
Sturtian Ice-Cap — for Ice-Cap it must have been — actually rested for a long" 
period of its existence upon the rocks that underlie its glacigenc- depositions. In 
these areas, as the ice finally receded, a mantle of greater or less thickness of 
glacial and fluviogiacial debris was spread over the glacially eroded surface of 
the pre-existing formations. In other areas deposition took place in deep water. 

The old highland areas existing during that glacial epoch, such as in the 
neighbourhood of Olary, the Barrier Rangers, Eyre Peninsula and in the vicinitv 
of Mount Painter were alt ice-capped and shed glacial debris into the sinking 
geosynclinal area. Even portion of the latter carried a capping of ice, so that 
earlier sediments of the Adelaide Series were in some areas consuierablv envied 
by the ice sheet before the glacial deposits Mere eventually laid down over their 
ice-eroded surface. 

During 1 he past 40 years the author has made extensive field observations 
of evidence bearing on problems concerning this period of glacialion in Australia. 
'1'he broader findings are as just mentioned, but further elaboration is reserved 
for a more detailed account in preparation. 

Lamixated (Tapley Hill) Post - Glacial Fo;o,iAtio>/ 

Following- stmigraphienlly above such beds of uncpjesrionable giaeigene 
origin there is always a thick formation of shale, or its slate equivalent, con- 
stituted of thin laminae which are not true varves, but the author is convinced 
do represent a modified type of seasonal deposition. This conclusion was reached 
long ago and voiced before the geological section of the Australian Association 
for the Advancement of Science at its meeting in Adelaide in January, 1907. 
Since then further convincing proof has been accumulated, and is bving dealt 
with in the compilation already mentioned. 

Tbe ribbon slates of this horizon in HowclnVs Adelaide Series have Jong 
been distinguished as the Tapley Mill Slates or shales as the case may he. In 
the main, they represent rock floiir washed out from the glaciated land during 
the waning phase of the glacial period. 

In its upward extension the Tapley J Till formation includes more and more 
calcareous contributions. Eventually, in the upward sequence, some beds appear 
of the composition of argillaceous limestone, to he quickly followed bv a greater 
or less thickness of purer limestones of the Brighton Limestone horizon. 

At Mundalho Creek this section of sediments amounts to only about 650 feet 
in thickness, but m the vicinity of Copley it is much greater, namely, 3,598 feet. 
This is evidence that die neighbourhood of Copley was situated further from 

the old shore line. 

W Not to be co)t(u.sed with the Aronna Range, which is j>art of the main Klindten* 
Ranges and extends north from Wilntwa Pound, 


The Brighton Limestonk Hokjzon 

The limestones and associated argillaceous beds of this horizon are com- 
paratively shallow water depositions, for bands of intraformationaJ chip and pellet 
breccias appear in this portion of the sequence. The purer limestone formations 
frequently exhibit good evidence of. their biostromic character. 

At Mundallio Creek the main limestone formation of this horizon docs not 
exceed 54 feet in thickness. At Copley it is thicker. And Arknackous Sedimknts above the Brighton Limestone 

At Mundallio Creek, as elsewhere in :he State, the sediments of this section 
are mainly of a reddish tinge of colour, often chocolate lo purple. 

At Mundallio Creek the measured thickness amounts (o only about 2,130 feet. 
According to Segnit (1939) these beds are much thicker than wc found them to 
be. We have found a vastly greater thickness further to the east in the deeper 
portions of the basim No complete section of this portion of the sequence is 
available for measurement at Cop]ey ? but to the south-west of Copley such beds 
are exposed on a major scale between Mount Scott and Mount Deception. Thus 
again supporting the contention (hat Copley is further east from the old shore 
line than is Quorn. 

Thk; Pound QuARtzmr (Cambrian) 

As the outcrops met with in the Quorn area are faulted and considerably 
eroded, our determination of the thickness, of this formation as met with there 
is subject to some revision, but it. is believed to be approximately correct. Our 
figure is about 2,000 feet. We cannot agree with Scgnit's 739 feet (1939). Also, 
we are sure that Howchin's figure of 800 feet for this quartzitc as estimated by 
him at Devil's Peak does not correspond to the true thickness in the Quorn area. 
The fact is that Devil's Peak represents only a badly faulted and sheared frag- 
ment of the Pound Formation. 

In the neighbourhood of Copley this horizon is developed in great strength 
upwards of 2.000 feet in the Mount Scott Range. 

At some 25 miles to the south-west of Copley, Sprigg- (1947) made an 
important discovery of fossils in a sandstone below the Archaeocyaihiuac lime- 
Mnne. This sandstone appears to helong to the Pound Formation, and the dis- 
covery lends added support to the contention that the latter is Cambrian. More 
recently we have visited the locality of this fossil find and have been able to collect 
additional specimens both of Sprtgg's medusa-like forms and of a plant-like 


In the South Flinders Ranges there has come Under our notice two other 
areas occupied by outcrops of this same quartzite, basal to the sedimentary 
succession of the Late Proterozolc geosynelinal basin. One is that traversed by 
.♦he Port Germein Gorge road, aJid the other constitutes the main mass of Mount 
Remarkable. The geographical situation of the three areas mentioned in the 
South Fmidcrs Ranges and of the Mount Aroona belt further north is illustrated 
m fig. 1. 

This Gkkat Quartzitk ok Port Germein Gorge 

The Port Germein Gorge road cuts across this quartzite at the northern end 
Of ?« north-south directed and northward pitching anticline. The conformable. 


dolomite -magnesite scries follows normally around the anticlinal structure; 
additional proof that this great quartzite is strati graphically below the magnesue 
scries as at Muadalliu Creek and at Copley and not just blocks of Pound 
Quartzite faulted into then present position. A portion of The Dolomue- 
Magnesite formation, trending in a general north-south direction, k to be seen 
in the bank of the Gorge Creek at the western entrance to the Gorge. Just a little 
farther to the west is located the Tertiary fault scarp of the Range; this effec- 
tively drops the Procanibrtan formations out of sight beneath the plain. 

Proceeding eastward through the Gorge where the highway is hewn 
through a trreat thickness of quartzite, a tuntier re-appearance of that of the 
Emeroo Range quartzite, the strike is observed to swing around from 
south-noith on the west side through west-east in the centre region to 
roughly north-south on the east side where the road emerges Ironi the quartzite. 
A little further on the magnesite series, having followed around the margin of 
the quartzite, again makes its appearance: here it is being worked by the "Broken 
Hill Proprietary Company, Still further on to the east, across the strike, and 
just beyond the locality Bangor, the Sturliati Tillite comes into view. 

Returning to the dolomite formation, it extends sonrh along the eastern Hank 
of the great quartzite anticline through the Wirrahara Forest and Beetaloo Water- 
works Reserve towards Crystal Brook. In that vicinuy also the tillite has been 
recorded (Howchin 1929). On its western side and along: its- southern end 1ho 
great quartzite is truncated by major faults of Kosciuskan omgeny. 

The Massive Quartzite ok Mount Rlmarkable 
Tfowchin (1916) was struck with the great thickness of the imposing ridgr 
of quartzite which constitutes Mount Remarkable. He suggested many possible 
ways of accounting for the great thickness; with none of which, however, was 
he satisfied. He had not conceived the existence in South Australia of a single 
quartzite formation of such magnitude. On its eastern side the country is thrown 
down by a great fault, again apparently of the Kosciuskan epoch. On its western 
side there appears to be some faulting also, but from the overlying Sturtian Tillite 
horizon there appears to be a regular upward succession to the west, culminating 
in a large development of another major quartzite formation which, judging by 
its broader features, appears to be the Pound Quartzite. In its relation to 
associated beds, there seems to he no doubt that the quartzite of Mount Remark- 
able is yet another outcrop of the great basal formation tinder consideration. 


Investigation of the several areas in the western Flinders Ranges discussed 
iu preceding pages leaves no doubt that in thai region, stratigraphically situated 
considerably below the record of the Sturtian glaeiatiou, and immediately below 
an extensive series of beds peculiarly rich in dolomite and magnesite. there exists, 
as the basal member of a very notable .section of later Proterozuic sediments, an 
extraordinarily thick development of quartzite, which is arkosie in part and 
subject to a great development of coarse conglomerate at its- base. In the Emeroo 
Range the lower section of this remarkable formation is exposed to view and 
is observed to exhibit a brown to chocolate appearance. There, sandwiched 
between some of the rudaceous and arenaceous beds are chocolate to purple shales 
as bands up to several feet in thickness 

The massiveness ot this formation is apt to cause it to h? confused with the. 
other extraordinarily thick arenaceous formation of that region— the Pound 


(Jusrtzite, The confusion is increased owing to the existence of purple shales 
and sandstone at the base of the latter as well as being found in association with 
the former. Fossils have never been observed in the older formation, but in the 
Pound Quartzite worm burrows are not infrequent, Recently Sprigg (1947) 
has made a very important additional fossil find, 


The Tent Hill Formation 

Segnit (1939) has. referred to the horizon of the Pound Quartzite., that 
extensive development of sandstones and associated conglomerates widely 
sprend over the neighbouring region on the western side of the Port Augusta- 
Lake Turrens Sunkland. To the earlier geologists who have discussed the 
geology of the region west of Port Augusta, the age of this arenaceous formation 
has been an intriguing problem. The formation is usually met with in the form 
of widely scattered mesas and bttttcs, obviously remnants of a former grand 
developmenl. The descriptive term, Tent T Till Formation, has long been applied 
lo it. This term came into common Usage from the resemblance in profile of 
some of the outliers or residual hills of the formation to that of a marquee tent. 

In the absence of fossils it has been the custom in the earlier records of the 
Geological Survey Department to doubtfully refer the Tent Hill Formation to 
the Ordovician. In the more southerly portion of the region, Dr. Jack (1914) 
has referred to several outcrops which may be considered as representing phases 
of this formation, including conglomerates and quartz.ites of the Corunna Range 
near Iron Knob, the quartzite of the Blue Range, in the Hundred of Verran, the 
Moonaby Range conglomerate. 

In this same general area, Segnit (1939) refers to this horizon the 
extensive plateau formation in the neighbourhood of Lincoln Gap, atid the rocks 
of the Comma Range, Mount Laura, Mount VVhyalla. Mount Young and 
Hummock Hill. All arc referred by him, as earlier mentioned, to the Pound 
Quartzite which is his D7 formation, which, by the way, is considered by him 
to be the topmost horizon of the Preeumhrian. 

The formations under consideration in all these localities are conglomerates 
and quartzitcs, usually flat -bedded or hut little inclined, not obviously meta- 
morphosed and with no indubitable signs of great antiquity, All repose with 
evident unconformity upon highly metamorphosed sediments and granites. Fig. 5 
is a section illustrating the relation of the Tent 1-iill formation of the western 
plateau region to the block-uplifted folded rocks of the South Flinders Ranges. 


T £S e 


: S' 







" "-?£**£** 

I*ig. 5 

liiK Ctmu-XKA Rangi-: Rvdacixiks and Akenaceous Formation 

We have examined in some detail an example of this formation as occurring 

in the Corunna Range,, located nnrrh of and within several miles of Iron Knob 

iownslnp Tins locality is well illustrated in pi. xiv. 

In the neighbourhood of the reservoir at Corunna Range, the bedr &lHlrtj 

N. 35° \V. and dip to the north-east. At the base of the series the dip Ls as 

steep as 30°i but rapidly diminishes in the upper beds to the east. Actually the 

Range is the remnant of a synclinal basin, for on its eastern side the dip is 

reversed, directed at low angles lo the west 


Ar the base arc red arenaceous and argillaceous beds; then follows a remark- 
able development ot conglomerate with some bands of quartz, gnt. A lower 
100 feet thick of greyish, medium-coarse, arkosic grit Is followed in vertical 
succession by a whiter phase containing larger pebble? Pacing upwards another 
100 feet an extremely coarse conglomerate is encountered. Here masses of the 
rock are composed of boulders of 4 inches to 6 inches diameter; occasional 
boulders were met up to 15 inches long, Current-bedding is evidenced. 

Most of the boulders are ot milky quart/., while occasionally are met 
examples of quartzite rich in hematite of a kind exactly resembling the qtiartzite 
of the Iron Monarch Hill Red jasper pebbles tip to 10 inches diameter are of 
rrcquent occurrence. Several examples ot quartz-museoviUvschist and banded 
jaspdites were met with, Occasional pebbles ot typical Gawler Range porphyry 
were encountered. At some horizons there appear in the fine base of the con- 
glomerate spangles and grains of pure hematite. 

No evidence was forthcoming to indicate that these boulder beds were in 
any way associated with glaciating The total thickness of this foniuitiim as 
exposed jn the Cortmua Range is said by Segnit (1939) to be at least 350 tect. 

Dr. Jack (1922) was the first to record finding in this conglomerate pebbles 
of feldspar-porphyry indistinguishable from that of the Gawler Ranges Oying 
to rhe west), also" of ferruginous jaspih'te and hematite similar to the fonnatiou 
:it Iron Knob and Iron Monarch (located a few miles to the south), jack states 
that at about 11, miles north-north-west of the Monarch is the easternmost 
exposure of the great feldspar-porphyry mass that forms the Gawler Ranges. 
Also relevant to our investigation is Eel wards' ( [936) record that pebbles of the 
ancient Middleback terrain are found in the conglomerate of the Moonahy Kantrc. 

Tim? ii is quite clear, from this evidence alone, that the Tent Hill Forma- 
tion Js younger than cither the Gawler Range Porphyiy or the Iron Knob terrain. 

Kouivalmko: or wte Coruxica Rakot. Fokmateon with that of thk 

F.weuoo Ran tu : 

The latest evidence bearing upon the question of age is our discovery that 
the lower 800 leer at least of the Kmeroo Range formation is composed of con- 
glomerates, quartzites and red shales, pctrologically similar to counterparts in the 
Corunna Range. No such association has been found in any outcrops of the 
Found Formation. Further, the range is strikingly similar to that of the con- 
glomerates of the Corunna Range, so similar that this fact alone is most con- 
vincing as to the identity of these two. 

It thus becomes clear, as already recorded (Mawson 194$}, tha. the Flinders 
Ranges east of Fort Augusta (see pi. xv\ fig-. 2) represent the crumbled sediments 
of tin. Late Proterozoic to Cambrian geosynclinal basin, while the Tent FIdJ 
Formation to the west of Port Augusta (see pi. xv, tig. 1) represents shallow 
na&tf sediments of the early stages of marine transgression over the coastal fringe 
of Yilgarnia, and corresponds to the basal formation of the Adelaide Series. 

A further important deduction that arises is that we can now fix lite a^e ol 
the Gawler Range Porphvry as pre-datirig the base of the Adelaide Scries. _ It 
may be tentatively regarded as a product of the period of Algoman revolution 
{ U.S.A. nomenclature ) - 


While the succession on the western side of the basin in the region of the 
Flinders Ranges can now, with a high degree of confidence, be correlated with 


that in the neighbourhood of Adelaide, the relation between the sediment* 
♦lipping to the east on the eastern side of the Mount Lofty Ranges and those of 
the Adelaide Series as established on the west side of the Range smd ia the 
Flinders Ranges is not so evident. 

IJowchin held that the sediments on the cast side of the Mount Lolry Ranges 
which dip consistently to the east and become successively mote metamorphosed 
as they progress in that direction are Ihe same beds as constitute the scries, the 
sequence of which he had established on the western slopes; the succession being 
repeated on Ihe eastern side of the Rarossian core of Ihe Raiiqc. Knr instance, 
he regarded the Grey Spur conglomerate, which unconfonnably overlies Barossian 
gneisses and faces east, as the Base of the Adelaide Series and equivalent to the 
Aldgate sandstone. 

We have since traced the Grey Spur conglomerate, to murk-fellows Creek, 
near Mount Magnificent. Accordingly, on II o\y chin's reckoning, the Black- 
fellows C'rcelc conglomerate and the overlyiug Mount Magnificent Series of beds 
should be the basal member oi the Adelaide Series. However, as already stated 
{Mawson 1939), the marble in that acnes is almost free from magnesia, and is 
therefore unlikely to be the counterpart of the richly magnetic dolomites of the 
lower Adelaide Series as developed on the neighbouring western side of tjhe 

Consequently it is obvious that, before Flowchin's views regarding a repeti- 
tion of the Adelaide Series on the east side of the rnnge can be accepted without 
further question, a thorough-going investigation must be made, Hossfeld (1935) 
holds that in the Wilhamstown area and to the east I hereof, the Proterozoic rocks 
belong to au older series (his Para Series) than those (his Narcoota Series) 
developed Sd well on the western flank of the Mount Lofty Ranges. 

There is considerable evidence to support the contention that much of the 
eastern flank of the Mount Lofty Ranges is occupied by rocks of Mosquito Creek 
Age. Certainly in the Olary - Broken Ilill region there is a large development of 
.sedimentary rocks intruded by the younger little-stressed granites, which underlie 
with violent unconformity the Stunian rillite formation and may be regarded as 
vslratigraphically equivalent to the Mosquito Creek Series of Western Australia. 
These intrusive granites have associated with them pegmatite apophyses notable 
$or containing beryl, tan tali tc, cnlumbite and uraniferous minerals, just as in the 
case in Western Australia. 

Thus the stage now reached U that, henceforth, we shall regard the Mount 
Aroona quartzitc as equivalent to the base of the Adelaide Series which, in the 
vicinity of Adelaide, is taken to be the Aldgate Sandstone. But the equivalence 
of ihe (jrcy Spur- Blackfellows Creek basal boulder beds and the Mount Aroona 
qnartzite, while possibly correct, has yet to be convincingly proved. 


The Late Proterozoic, Nullaginc Series, of the north-western and northern 
Australia has long been correlated (Cotton 1930) with the Adelaide Series. In 
Central Australia the Peitakntirra and Pertatataka Series exhibit a close relation- 
ship in sequence of beds with the exception that the glacial horizon has not yet 
been recognised there. 

As already published (Mawson 1946), a comparison with the Prc-Cambrian 
record of South Africa is convincing that this division of the Australian record 
has its equivalent in the Transvaal system and extensions of the latter in other 
areas of African Hie detail of the South African succession bears a "broad 


general correspondence to that existing in South Australia. Thus the Daksport 
Glacial Horizon, the Lower Griquatown TilHte and the Numees Tillite. (all 
apparently records of the same glacial epoch) may be said to be broadly equiva- 
lent to our Sturtian Tillite. The underlying dolomites and overlying limestones 
rich in 'algal" structures are a feature of the sequence in both case^. The gieat 
conglomerate-quarlzite formation at the base of the Transvaal System, resting 
with violent unconformity upon older formations, appears to correspond to the 
conglomerate -quartzitc formation of the Emeroo Range and the Tent Hill Forma- 
tion ol Yilgarnia further west. 


Consideration is given to ihe succession of sediments occupying the great 
gcosynclinal basin which developed olT the eastern margin of Yilgarnia during 
Late" Proterozoic to Middle Cambrian time. The present contribution deals only 
with accumulations along the western side of the basin in the region of the 
Flinders Ranges. Subsequent comributions will deal with depositions respec- 
tively of the eastern margin and of the central region of the depression. 

The sediments accumulated in two locations near the western side of the 
basin are considered in some detail. The first is the neighbourhood of Copley, 
the second is the district to the west of Quorti. The latter is relatively nearer 
to the western margin of the gcosynclinal basin than in the former. 

The sediments considered are of Upper Proterozoic age, equivalents ot the 
Adelaide Series of the Mount Lofty Ranges and of the Nuilagine Series of 
North Western Australia. 

Attention is drawn to a remarkable correspondence, so far as the broader 
features of sedimentation of this geological period are conrerned, of the Aus- 
tralian record with that of South and Equatorial Africa. 

Finally, the basal formation 01 the sedimentary accumulation in this gco- 
synclinal area has been traced westward as an attenuated transgressional deposi- 
tion overlapping the margin of Yilgarnia where it is known as the Tent Hill 
Formation. This settles the age of this latter feature, a long-disputed problem 
in South Australian stratigraphy. 

A further deduction of some importance is that both the Gawler Range 
Porphyry and the iron Knob iron-ore formation pre-date the deposition of the 
Adelaide" Series. Reference is made to the occurrence in the Olary - Broken Hill 
region of sediments of Mosquito Creek age underlying the Sturtian glacial 
horizon, and it is suggested that further derailed field investigation of the sedi- 
ments and meta-sediments of the eastern flank of the Mount Lofty Ranges may 
prove them to be of similar age. 


The photograph from the summit of Iron Knob is reproduced by courtesy 
of the Broken Hill Proprietary Coy. Tn the prosecution of field work associated 
with this contribution, I have been assisted by the co-operation of students, 
amongst whom H. E. E. Brock and K- C Sprigg have been notably helpful. 

Andrews, E. C 1922 "The Geology of the Broken Hill District." Memoir* 

of the Geol. Surv. N.S.W., Geology, No. 8 
Andrews.. E. C. 1937 "The Structural History ot Australia during the 

Palaeozoic. " Proc. Roy. Soc. N.S.W., 71 


Barnes, T. A... and Kleeman, A. W. 1934 "The Blue Metal Limestone and 
its Associated Beds." Trans. Roy. Soc. S. Aust., 58 

Bryan, W- II- 1932 "Early Palaeozoic Earth Movements in Australia." 
A.N.Z, Ass. Adv. Science, 21 

Cottom, L. A. 1930 "An Outline and suggested Correlation of the Pre- 
Cambrian Formations of Australia." Pres. Add. Roy, Soc. N.S.W. 

Dickinson, S. B. 1942 "The Structural Control of Ore Deposition in some 
South Australian Copper Fields." Geol Surv. S.A., Bull. 20 

Edwakds, A. B. 1936 "The Iron Ores of the Middlfhack Range. South Aus- 
tralia." Proc, Aust. Inst. M.M.. New Series, No. 102 

PTossfeld, I\ S. 1935 "Geoloj^ of Part of the North Mount Lofty Ranges." 
Trans. Roy. Soc. S. Aust., 59 

Howchtn, W. 1916 "The Geology of Mount Remarkable." Trans. Roy. Soc, 
S. Aust., 40 

Howchin, W. 1926 "Geology of the Barossa Ranges and Neighbourhood in 
relation to the Geological Axis of the Country." Trans. Roy. Soc. 
S. Aust, 50 

Howchtn, W. 1928 "The Sturtian Tillite and Associated Beds on the 
Western Scarp of ihe Southern Flinders Ranges/* Trans. Roy.. Soc. 
S. Aust., 52 

Jack, R. L. 1914 "The Geology of the County of Jcrvois and portion of the 
Counties of Buxton and York, with special reference to Underground 
Water Supplies." Geol. Surv. S. Aust., Bull 3 

Jack, R. L. 1922 "The Iron Ore Resources of South Australia." Geol. Surv. 
S. Aust., Bull. 9 

MAtncAK, C T. 1927 "Geology of the Wiliunga Scarp." Trans. Roy. Soc. 
S. Anst. 51 

Mawson, D. 1912 "Geological Investigations in the Broken Hill Area." Me- 
moirs Roy. Soc. S. Aust, 2, pt. iv 

Mawson, D. 19.^9 "The First Stages of the Adelaide Series: As illustrated 
at Mount Magnificent." Trans. Roy. Soc. S. Aust., 63 

Mawsox. D. 1941 "Middle Protcrozok Sediments in Lhe Neighbourhood of 
Copley.'* Trans. Roy. Soc. S. Aust., 65 

Mawson. R 1942 "The Structural Characters of the Flinders Ganges. H 
Trans. Roy. Soc. S. Aust.. 66 

Mawsox, D. 1946 "The Geological Background of South Australia." Hand- 
book of 25lh Meeting of A.N.Z.A.A.S., Adelaide 

Seonit, R. W. 1929 "Geological Notes from the Hundred of Adams. Flin- 
ders Ranges." Trans. Roy. Soc. S. Aust., 53 

Segnit, R. W. 1936 "The Geology of the Northern Part of the Hundred of 

Macclesfield." Geol. Surv. S. Aust, Bull. 16 


Segnit, R. W. 1939 "The Pre-Cambrian - Cambrian Succession." Geol. 
Surv. S. Aust., Bull. 18 

Sprigg, R. C. 1942 "Geology of the Eden - Moana Fault Block." Trans. 
Roy Soc. S. Aust., 66 

Sprigg, R. C. 1946 "Reconnaissance Geological Survey of Portion of the 
Western Escarpment of the Mount Lofty Ranges." Trans. Roy. Soc. 
S. Aust., 70 

Sprigg, R. C. 1947 "Fossils from a Sub-Cambrian Horizon in South Aus- 
tralia." Trans. Roy. Soc. S. Aust., 71 

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Vjtvv "i the Lincoln Clap Range fottiTi the mrpu higltwaj ta the w**t tti 
fori Auuu-tit. This i- a iorniatfcin trf fun slight b inclined rtfUrtzites and 

pi ,',:, betU Ii :<1H.,uiN "m nw-l.!|, ,n fch'p l)a<al !>).-iriation of Jm 

a ,i, laSrjo s-ui tuitu the hMil maw Pi Yilgarirta. 

Tin- tyactrrn scarp nice ol tli-c K ■■" Ki,, ui' as \i>.'\vci trotn the Port 

Aufcii&tu Lafei' '-1 wrens siinklaiiD Hie cautc raiiRe it ititutod i>1 

steeply dipping berl* of tire nuhiiTnus ami arenaceous hasal formation o] the 
Mvftidi' Slim.- and represent 1 ; deposition? along flic wptferd margin 

, .1 Ulf PlVtr fl I OS} rtcHiWi 


ByT. Langford-Smith 

The survey includes an area of over a hundred square miles in the Jamestown district. The beds 
consist of Proterozoic sediments, which have been folded into synclines and anticlines with north- 
south axes. Jamestown is situated in an anticline which has been differentially eroded to produce a 
series of parallel ridges and valleys, while to the east of the town is a broad synclinal valley. 


By T. Laxci-okd-Smitk * 

[Read 9 October 1947] 


Summary- * ..281 

Introduction , , . . , , „ _ 281 

Details of Stratigraphy t , _ 284 

Faulting and Folding . . . . . . . . ., . . , . . , . . 295 

Acknowledgments .. , , ., , , tI 295 

Rfj-t.rencf.s .,295 


The survey includes an urea of over a hundred square miles m the James- 
town district. 

The beds consist of Proterozoic sediments, which have been folded into 
synclines and anticlines with north-south axes. Jamestown is situated in an anti- 
cline which has been differentially eroded to produce a series of parallel ridges 
and valleys, while to the east of the lown is a broad synclinal valley. 

There is a fair degree of correlation between the sediments in the vicinity 
of Jamestown district and the Precambrian beds in the Adelaide district, the 
vertical range including the beds between the Upper Phyllites and the Brighton 
Limestones of Howchin's classification. There are also very marked correlations 
between the Jamestown beds and the Precambrian al Appila Gorge and Orroroo. 

The sediments below the glacial beds are about 13,700 feet in thickness. 
They include phyllites, slates, hornfcls, shales, quartzites, dolomites and dolomitic 
limestones. A feature of this group is the large number of thin bands of dolomitic 
limestone, Phyllites and slates dominate the lower portions. In the central 
section is a bed of arkosc sandstone and quartzite about 200 ieai thick. Above 
the latter are more slates and phyllites, and also two distinctive beds of quartzire, 
the upper of which underlies the glacial beds. These beds of quartzite outcrop 
very prominently in the highest points of the Campbell Range, and its northern 
continuation through Mount Lock. 

The glacial beds are noted for their great thickness (3,000 feet), They 
include a variety of tillite and fluvio-glacial sediments. Erratics are numerous. 

Above the glacial beds is a transitional zone (300 feet) of fluvio-glacial 
sediments, banded slates, and thin bed of limestone. Above this occur about 
10,000 feet of shales and slates with the typical Taplcy Hill ribbon banding, 
which in turn are overlain by a series of banded siliceous limestones. 

The. field work of the following paper was carried out in 1941 and 1942, 
while the writer was a member of the Council for Scientific and Industrial 
Research party engaged on a Soil, Soil F.rosion and Tarid Use Survey of County 
Victoria (fig. 1). It is the first of a series of papers dealing with various 
aspects of the Geology and Geography of the County Victoria area, publication 
of which was delayed on account of the war. 

* Department of Post-War Reconstruction, Canberra, A.C.T, 
Trans. Hoy. Soc. S. Aust., 71. 12), 1 December, 1917 


J*$MJjtrzfi Maef*'? 3 ^* 

Fig. t 





Banded siliceous limestones below 
Brighton limestones, or basal beds of 
Brfghton limestones 

T«pley li'H ribbon slat* 

Sturtian fill ite 

Sub- glacial quart zite 

Glen Osmond upper slates 

Miteham and Glen Osmond quart zite 

Glen Osmond lower slates/blue metal* 
limestones, upper phyllites 




Banded iikeous limestones 

Tipley Hill series 

° Tillite scries 


o Sub -glacial quartzites 

o Slates, phylfites and hornfels 

$, Quartzite and sandstone 

Slates and phylfites with thin bed* 
of dolomilic limestone 

Quartzites and sandstones 

§ Phyllites and slates with thin beds 
of dolomitic limestone 

26,700 FEET „. 

Fig. 2 


In the neighbourhood of Jamestown very little geological work has been 
attempted in the past. Investigations have been carried out by liowchin and 
Segnit in the Appila Gorge near Tarcowie, by Howchin in the Orroroo district, 
and by Hossfeld in the North Mount Lofty Ranges. The detailed stratigraphy 
of the immediate vicinity of Jamestown has been neglected. r\o doubt this has 
been due largely to the scarcity of surface outcrops in this district. 

In 1941 the complete coverage of the district by air photographs of a scale 
4 inches to 1 mile made a new approach possible, as small, obscure outcrops 
were dearly shown in numerous localities. Apart from natural rocky gullies, 
soil erosion gullies of recent occurrence proved of great assistance, as they 
frequently exposed the bed-rock for long distances. Scattered through the 
ploughed fields, numerous low rocky outcrops which otherwise would have been 
most difficult to detect were revealed by the photographs. Not only were the 
photographs invaluable in the detection of outcrops, but also in later mapping 
work. Outcropping series could be accurately traced for miles with great 

In general, the region consists of parallel alternating ridges and valleys, 
running in an approximate north-south direction. The area covered by the 
survey is folded into a syncline to the cast, and an anticline to the west. The 
syncliue conforms in topography to a broad, gently sloping valley, while the anti- 
cline (on which Jamestown is situated) has been differentially eroded to form 
an alternating series of ridges and valleys. 

As a result of the investigations, it is now evident that many of the beds 
in the vicinity of Jamestown are largely analogous to beds in the Adelaide district 
described by Howchin and others. The vertical range appears to correspond 
to Precambrian beds of the Adelaide district between the Upper Phyllites and 
the Brighton Limestones (fig. 2), 

Sections were run eastwards from the tilHte-contaming ridge directly south 
of Mount Lock in the Hundred of Caltowie. to the tillite-containing ridge of the 
Browne Hill Range in the western part of the Hundred of Whyte, a distance of 
about 10 miles. These sections are shown on the accompanying plan and in fig. 3 
by the lines PQ and RS. PQ runs in an easterly direction six miles north of 
Jamestown, and RS continues the section along a path running almost due east 
of the town. 

Considerable work was carried out both north and south of the section lines, 
and eventually an area of over a hundred square miles was surveyed and mapped 
in detail. The findings of the survey are presented in the following text, with 
accompanying sections, maps and photographs. 



This series is in the form of a large eroded anticline which has been pro- 
duced by folding through a north-south axis (see section * 4 PQ"). It is bounded 
to both east and west by quartzite and sandstone ridges (**B'* and "Bl"). In 
the few exposures where the junction between the series "A" and il B and Bl"' 
could he observed, the rocks of "A" were contorted, and it was not possible tn 
determine whether there was any unconformity. From the general trend of the 
dips, however, it was apparent that if an unconformity did exist, it was not 
particularly marked. 




Ski 109 

Dips were, measured at numerous localities through series U A". They 
varied from about 45° W. near "M" through the vertical to about 70" & near 
U R". Except for some slight local variation due to minor folding the gradation 
in dips through the range of angles given was comparatively regular. 

The width of outcrop of the beds of this scries ranges from 4,000 feet in 
Sections 140 and 139, Hundred of Belalie, to 9,300 feet one mile north of the 
Reialic - Mamiauarie Hundred line. Compression has badly crushed and folded 
the former region, and the areas further south near Jamestown arc almost com- 
pletely covered by soil; thus detailed investigation was restricted to the northern 
areas' towards Mount Lock. Here the true thickness of the beds is approxi- 
mately 5,000 feet. 

The rocks of the series are highly metamorphosed. Thyllites easily dominate 
the group, hut slates, some of which are arenaceous, are fairly well represented. 
Pyrites crystals in the form of little cubes are quite commonly found in both 
tbesc types; they frequently show decomposition to limonite, and often cubes of 
limonite occur as pseudomorphs. Both phyllitcs and slates, when fresh, arc 
characirxistically grey to bhte-grev, hut show a great variety of colours when 
weathered. Shades of yellow and red as a result of iron-oxide staining are 

A few thin bands of quartzitc impregnated with quartz veins were found in 
parts, but this was not typical of tbe series. 

A number of very thin beds of dolomite and dolomitic limestone occur 
through the series. These are well exposed along the road to the west of Sections 
156 and 157, Hundred of Belalie, where they dip nearly vertically. A sample 
of these dolomitic limestones was subjected to a rough quantitative analysis, the 
assumption being made that calcium and magnesium would be almost entirely 


in the form of carbonates, and iron and alumina of sesqui-oxides. The following 
figures were obtained : 

SiO„ - 24% 

CaCO, - - - 45% 

MgCO a - 20% 

Fe,0, 4 ALO, - - 6% 


Ilowchm (1) described as "Glen Osmond Lower Slates" beds in the Orroroo 
district which closely resemble the series "A". He found calcareous zones in 
these slates, which "he believed represented rhc Blue Metal Limestones of the 
Adelaide district. 

Mawson (2) has compiled a detailed description of Middle Proterozoic beds 
underlying the tillite in the Copley district. He found a large amount of dolomite 
in these beds, which appears very similar to that in the series "A". 

Hossfeld (3), in his surveys of the North Mount Lofty Ranges, defined two 
distinct series, the Para (older Adelaide) and Narcoota (younger Adelaide), 
which were separated by an unconformity. He assumed that the" basal beds of 
the Narcoota Series were contemporaneous with Howchin's Mitcham and Glen 
Osmond Quartzites (4). There is little doubt that the quartzites "B" and "Bl" 
in the Jamestown district are contemporaneous with the Mitcham and Glen 
Osmond Quartzites, and it would therefore appear that they are representative 
of Hossf eld's basal Narcoota beds. Although there is no proof of unconformitv 
between beds ''A", "B", and "El", it is considered that the series "A" must 
represent the upper beds of the Para series. 

Summing up, it is most probable that the series "A" is contemporaneous on 
the one hand with Howchin's Upper Phyllites, Blue Metal Limestones, and Glen 
Osmond Lower Slates of the Adelaide district, and on the other with Hossfeld's 
Para series in the North Mount Lofty Ranges. It also appears similar to Middle 
Proterozoic beds described by Mawson at Copley, and by 1-Iowchin at Orroroo. 


"TT and "El" are respectively the easterly and westerly outcrops of an 
anticlinal fold. 

The Eastern Outcrop ("B") 

The beds of "B" are exposed in the ridge which strikes roughly north and 
south through the central portions of the Hundred of Belalie. The direction of 
strike passes through Jamestown, although uo outcrop is visible in the town itself, 

Proceeding from the. town in a northerly direction, the scries is first exposed 
in a low ridge immediately west of the railway crossing, where it has been 
quarried. The ridge rises gently for about % miles, and then falls again, and 
finally dips out of sight in Section 8S, Hundred of Mannanarie, approximately 
8 miles north of Jamestown. 

Half a mile south of Jamestown there is a small outcrop in the form of u 
low hummock a quarter of a mile in length, where the rock has been quarried 
for road metal. A mile further south it outcrops in Section 75. Hundred of 
Belalie, and forms a gently rising ridge which continues in a southerly direction 
for several miles. 


The rocks of U B" are highly felspathic, the fdspar in most cases showing 
an advanced state oT decomposition, which gives the rock a very distinctive 
speckled appearance. In many instances the kaolin has been weathered away, 
leaving numerous small cavities between the silica grains, which causes the rock 
to crumble easily. Textural gradations within the series range from fine-drained 
sandstones and quartzites to grits. Sandstone is dominant, hut siliceous quartzitcs 
do occur. Quartz veins are common throughout the series. When these veins 
were found 'in the sandstone it was noted that they had not affected the impreg- 
nated rock, indicating the intrusion of cold siliceous solution. Well-defined ripple 
marks are a feature of the scries, implying shallow water deposition. 

At a distance of 44 miles north of Jamestown there has been some strike- 
faulting, resulting in repeating outcrops of the beds for a short distance, This 
faulting has been caused by the same pressure which was responsible for the 
faulted zone to the west (see plan). 

The true thickness of "B M ranges from 150 to 250 feel, and the beds dip to 
the east at 65° to 72°. There is no evidence of unconformiiy hetween "IS" and 
the overlying shales and slates of W C" 

The Western Outcrop (Rl) 

W B1" outcrops in a high ridge half a mile east of Mount T-nrk (sec plan) 
running almost due north and south of this point. 

To the north, thi* ridge was followed as far as the Mannanarie-Tarcowte 
toad, and from here it was seen to be continuing in a northerly direction. 

To the south the ridge was traced to Section 140. Hundred of Eclalic. From 
here to Section 41, Hundred of Caltowic. there has been a great deal of faulting 
and shattering. From the latter point the series again outcrops in a ridge, and 
this was followed (or five miles to the south, into the heart of the l:hmdaleer 
Forest. There has been much faulting in this area, and this has resulted in the 
repeated surface outcropping of "HI". Close to the eastern margin of one of 
these outcrops, located in the forest just west of Section 287, Hundred of Belalie. 
is a bed of crystalline limestone, composed of large calcite crystals. This bed 
was followed for some miles to the south, and was found to be the same as that 
mentioned by Lockhart Tack (51 in connection with the occurrence of rock 
phosphate in the Hundred of Reynolds. 

The beds of *&!" are almost identical in rock type to those of "B", although 
they arc generally more highly metamorphosed. Near Mount Lock the rock in 
general is a fine-grained, highly siliceous, and extremely hriitlc quartzite. It is 
often arkosc, although some of the more highly silicilied types do not appear to 
contain any appreciable felspar or kaolin. Very small pyrites crystals were 
observed in one specimen. 

Howehin (loc, cii.) has described some quartzile beds near Mucr;i Hilt as 
Glen Osmond Quartzites. Since. Mqcra liill is only 10 miles north of the outcrop 
of "Bl" on the Mannanarie- Tarcowic road, and also since "Bl" could be seen 
continuing in the form of a rid^e for some distance to the north, it is probable 
that the outcrop near Mucra Hill is a continuation of "HI". In any case, there 
is little doubt that "ft" and "Rl" represent the Mitcham and Glen Osmond 
quartzite of the Adelaide district. They would thus also be representative, of the 
basal beds of Hossfeld's Narcoota series. 


Slates A>m Pmvlljtes with some Siialks and Horxfels. 
ani> Tni.\ Bfcns ok Dor-nMrnc T-imestone 
The beds "C and ^Cl" overlie "fl" and "Kl'V and are conformable to them. 


This was clearly shown in Section 137, Hundred of Belalie, and in areas near 
Mount Lock. 

The Eastern Outcrop (C) 

The beds consist mainly of grey to dark blue-grey slates and phyllites, with 
some shales and grey hornfels. The slates are mostly thinly laminated, although 
there are a few instances of strong banding. Much of the rock is weathered on 
the surface to a soft, yellow-brown slate which shows the bedding planes very 
clearly. Cleavage planes are strongly developed and the cleavage angle is fairly 
consistent at about 65 C: to the west. The true dip of the beds raqgcg from 65° 
east near 'TV' to 75° east at "D". The average thickness is about 2,600 feet. 

The Western Outcrop ("CI") 

The rock types of i( Cl" are very similar to those of *£**. The slates in parts 
arc arenaceous, and there are a few thin bands of sandstone and quart/he, which 
sometimes occur as small lenses. There are also a number of very thin beds of 
dolomitic limestone. These outcrop in the eastern part of "CI" in the Mount 
T.ock area, in the bed of a small creek in Section 150, Hundred of Belalie, and 
on the road between Sections 120, 116E, and 117, Hundred of Belalie. Several 
bands of very similar rock were found at the northern corner of the Bundaleur 
Forest Reserve. Analysis of a sample from Section 150, Hundred of Belalie, 
yielded the following results : 

SiO., - 37-9% 

CaCO. - 48*0% 

MgCp. a - - - - 21-0% 

AUG., + Fe.XX - - - 3'0% 


This analysis was conducted on the same basis as that of the dolomitic lime- 
stone in "A". 

There is evidence of some minor folding and faulting in the beds of "Cl" 
just east of Mount Lock peak, and it is evident from the accompanying geological 
plan that there has been considerable pressure in this area. It is consequently 
difficult to measure the. true thickness of the beds. On the line PO (see plan) 
the thickness is approximately 3,500 to 4.000 feet. 

The beds of "C" and "CV* appear to represent the lower portion of the 
Glen Osmond Upper Slates in the Adelaide district. With the exception of the 
dolomitic limestones, they also appear analogous with the lower members of beds 
in The. Orroroo district described by Howchin (loc. cit.) as Glcil Osmond Upper 
Slates. i r 

SERIES (, D" AND "Dl" 

The Eastern Outcrop ("D") 

Tins is a prominent bed of quartzite aud sandstone, which, however, has an 
average thickness of only 18 feet. The rock is fine-grained, and is white to 
yellowish-brown in colour when fresh. When found as a quartzite it is sometimes 
very compact and brittle. The bed is impregnated with numerous quartz veins. 

Outcrops occur as a series of small hummocks. These were traced from 
Section 120, Hundred of Belalie, in a northerly direction, as far as Section 325. 
The dip was found to be fairly constant at 75°" to the cast 

A small quarry in Section 135, Hundred of Belalie, shows that beds "C" and 
"E", on either side, are conformable to "D". 


Tile Western Outcrop ("Dl") 

The beds of (, D1" comprise the peak of Mount Lock. Together with the 
sub-glacial quartzites "Fl'\ they constitute most of the highest parts of the range 
containing- Mount Lock. They also constitute the highest parts of the Campbell 
Range on the western margin of the Bundalecr Forest, the latter range being 
the southern continuation of that containing Mount Lock. Both i{ DV* and the 
sub-glacial quartzites "Fl" form the backbone of the two parallel ridges which 
are a very prominent feature of these ranges for many miles, to boih north and 
south of the Jamestown area. 

The thickness of '"Dl" is very difficult to measure on account of the large 
amount of rock talus on either side. At Mount Lock the thickness is between 
45 and 90 feet, and the dip 45° to 48° to the west. The beds "CI" and "HI" on 
either side appear to be conformable. The rock is very similar to that in "D", 
except that it is morely highly metamorphosed. Most of the quartzite is dense, 
siliceous, and very brittle, and contains quartz veins. It is extensively fractured' 

The quartzite "Dl" has its counterpart in the Appila Gorge/some miles 
north of the Jamestown area. In Fact, comparison between the outcrops in both 
places showed that the whole sequence of beds immediately below the tillite 
coresponded very closely. Segnit (6), from his observations in the Appila Gorge, 
concluded that a large part of the Middle Hrecambrian formations arc missing. 
and have either been faulted out by strike faulting, or removed before the deposi- 
tion of the tillite. If this be so, then the great similarity of the sequence in the 
Jamestown area infers that here also the Middle Precambrian beds must be 
missing. However, it seems to the writer most improbable that strike faulting 
should have occurred in exactly the same manner and position in these two 
localities. Also, if the missing- formations had been removed by erosion, then an 
identical amount of erosion must have occurred at both places, which seems very 
unlikely. It is not considered, therefore, that there, has been any large-scale 
removal of hedt, either here or at Appila. 

Slates, Piiylutes, and Hokkeels 

With the exception of the dolomitic limestones found in "CI", the beds o J 
[ *E" and "El" arc identical with those of '"C" and "CI", consisting- of slates auc 

phyllites with some hornfels and shales. The slates, in parts, are arenaceous. 

The beds of "R" are 1,800 feet thick, and dip at angles ranging from 
70° to 81 G to the east, * 

There is great variation in the apparent thickness of "Kl", as these beds 
have, in parts, been subjected to intense pressure. Near Mount Lock the thick- 
ness is about 600 feet. There has been considerable compression due to folding 
in the neighbourhood of Section 44, Hundred of Caltowie, while about 7 miles 
south of this point, in the Hundaleer Forest, the thickness increases to about 
1,000 feet. 

Near Mount Lock the dip is west at .aboul 47°. The beds arc conformable 
with those above and below ("DP and "Fl"). 

Sup-glacial Qoartzitks 
The EASTkK.v Outcrop ( 4J F") 

The sub-glacial quartzite iv F" which underlies the tillite in the central parts 
of the Hundred of P.elalie, is not a prominent physiographic feature, as ss com- 


monly the case elsewhere. It outcrops on the lower western slopes of the titlite- 
containing ridge '*</* In some areas it is almost entirely obscured by talus and 
soil, and as even the best exposures are not complete it was impossible to secure 
much detail. 

The total thickness of il F' is about 100 feet, but it was not possible to deter- 
mine whether the whole of this was quartzite. Surface materia! indicated thai 
there arc intercalated beds of shales and slates. The dip of "F" was difficult to 
measure accurately, but it is approximately the same (81° E) as the underlying 
shales and plates of "E'\ There is no apparent unconformity between "F" and 
the overlying tillite "G n . 

The quartzite is characteristically brown in colour. It is fairly compact and 
is moderately felspathic. 

The Wkstkrn- Outcrop ("Fl") 

The sub-glacial quartzites "Fl" are in two major beds, separated by beds 
of shales and slates, amongst which is a little tilhte. There is no tillite aL all 
below the lower quartzite. 

In the Mount Lock area each of the quartzite beds is between 20 and 40 feet 
in thickness, and the shales, slates, and tillites in between amount to about 130 feet. 

The only outcrop which provides a clear section through the series is about 
7 miles north of Mount Lock, in the gorge through which runs the Mannanark- 
Tarcowie road. The lower quartzite here is a massive bed 100 feet in thickness, 
while the upper bed is only 20 feet Ihick. Between the quartzite is 170 feet of 
shales and slates, which do not contain any tillite. The tillite commences abruptly 
above the upper quartzite, without any apparent unconformity. No unconformity 
could be detected between the lower quartzite and the beds underlying it ("El"). 
The Appila Gorge is only 6 miles due west of these outcrops, and the relation- 
ships between the exposures in the two localities are very close. At Appila the 
tillite commences very abruptly, resting on a bed of quartzite 50 feet thick. Cndcr 
this quartzite is about 15 feel of shales, and then another bed of quartzite 40 feu 
thick. It is notable that neither at Appila nor on the Mannanarie-Tarcowie roau 1 
is there any tillite between the two beds of quartzite, although a little tillite is 
present in the Mount Lock region in association with the shales mid slates. 

As previously noted, tbe sub-glacial quartzites "Fl" and the quartzitcs "D" 
form the central axis of the mountain range running north through Mount Lock, 
and continuing to the south as the Campbell Range on the western margin oi 
LUmcUdeer Forest. 

In Sections. 325 and 327, Hundred of Vangya, near the Bundalecr Forest. 
the sub-glacial quartzites and adjaeent series have been displaced by a consider- 
able dip fatdt, the horizontal displacement being about 2,400 feet, 

SERIES "G," "Gl." AND '"G2 ,! 
Thk Central Outcrop ("G") 

This rilbte outcrop forms quite a prominent ridge running north and south 
through the centre of the Hundred of Belalie. Except for the gorge near James- 
town, through which runs the Relalie Creek, the ridge constitutes a stream divide 
for many miles. Tbe average height of the ridge above the surrounding country 
is about 200 feet. 

The most useful section through *'G", or in fact through any of the tillite 
in the district, occurs in a small creek gorge running through the northern part 
of Section 303 and the southern part oi Section 160, Hundred of Belalie. about 
6 miles N.N.E. of Jamestown, The gorge has been formed by the headward 
























5HAL.L&, *I.*«C6 AND TIlLlftS 


fiwt ouvmed sAwiiaTomc: witm DAwE liLicincn quAimrt 


quartz ire 

shales - paotly msc^reo 



J*nos>chc v*irn humE«Oi>5 im,hi, rlaaCS Gr MustOvU 

HI Sfkft* 

SANDsroMfia an» TiLLiTaa. wrtlv 09it"M:0 


*"-OSt (AN Lift! ONE MIT Pi Q JAAT2 V{INI 




aoM okhh nAWrt 






n:n ".it* moh dh.ii 

>RO AM» SOFT ft i ...i . 

'« CR^Ancs 


•V"-.^? 6REy HaHWF€l_S [ -„ Q^itiiTI -£i!£ £ J CMS.- »i*My FLAf! -!(.£. »T PMlLiUHE 

!-.■'-•'-:■ .'.-"Jij IJNDilQrJL, Willi 5CH1F Qi_..»K ~ I I 7 C 

(JuART?,i tL 



TlvllTE WITH MAHV 5 SO Mf uf TO J'DlAWCTfft, » Nl^kr iM w 




till, re «vi: H nuwekous erratics u> m iicus. «>. diameter. 



S!;^ SE - i ? U . A,<Jr;£ ' r ^,^ h1 ^ lJ * (IT 2 w ElP»5, SOME CAVITIES IN QU*RTJ,TE MHWl 


riLi-iTE shows 4TKONG cowi'asswdh partly oescuafc 

S«avi<5 fROM QXIpd STAININ6 " ** ve.hj. 


s-u6'{,L*r.-A!. QLJ*«TZiT=: 

■ 1TI.Y oas;u(iej) 



erosion uf a -stream following the line of a slight dip fault which has caused a 
horizontal displacement of about 50 feet. 

It was Found that when continuous exposures of the scries could not he 
obtained from the sides and bed ol the creek, the nixing beds could very often 
be found outcropping: in the hillside above. Thus by the correlation of a number 
of small sections in the gorge, one almost continuous section was built up sbuwmg 
great detail. The main features of this section are illustrated in the accompanying 
d'a^ram (fig. 4). 

There is no evidence of major strike faulting in the series, ami the details 
id Lite section are such that if repetition of beds were present, it would almost 
certainly be revealed. There is little doubt that the measured section represents 
a continuous sequence. 

The most notable feature of the series is the great thickness of the tilJite. 
which amounts to approximately 3,000 feet. As far as the writer is aware, this 
occurence in the Tamestown district is the thickest Sturtian tdhte formation yet 
recorded Ir is considerably greater than that recorded by Scgnit (toc\ tfftj ni 
the Appila Gorge (1*586 feet). A recent summing up by Sprigs (/) oi work 
in the Mount Lofty Ranges indicates that the greatest measured thickness of 
fcJUitt there is about L.200 feet. 

The tillite series is underlain by the sub-glacial quarUites "F" (which have 
already been described), the tillite commencing very abruptly above die quartzile. 
Although unconformity has been recorded between the tillite and sub-glacial 
quartzite from other parts -of the State, there is no evidence of this in the James- 
town district. Unlike the base of the series, there is no abrupt upper hmit to the 
tillite A prominent feature of the upper part of the scries is a bed of very 
coarse arkose grit. It has a most distinctive appearance and proved most useful 
as- an index bed when examining the tillite outcrop "G2" on the eastern margin 
of the incline, in the Browne Hill Range. Above this arkose gnt the glacial 
and fluvuvglacial beds giadually grade into those of the Tapley Kill series. }h? 
rocks of the latter series exhibiting characteristic form from 250 to 300 feet 
ahuvc the grit. The transitional beds arc described in greater detail in the follow- 
ing section, M H". The fact that no sudden break is evident at the close of the 
glacial period would indicate that conditions gradually became warmer, introduc- 
ing a period when nuvio-glaeial deposition was dominant. Evidence from the 
J;imcstown area lends further support to rhe hypothesis that the Tapley Hill beds 
have a varve-like origin. 

There is less pure lillite in the upper than iu the central and lower parts of 
the lillite formations, Muvio-glacial shales and slates are dominant, and m many 
sections the only direct evidence of glacial conditions is the occasional presence 
of erratics. Although the central and lower horizons have a much higher pro- 
portion of pure tillite, even here intercalated beds of sandstones, quartzites, shales, 
and slates are very numerous. 

Erratics arc particularly numerous in parts of the series, and in some instance* 
range up to 3 feet in diameter. The erratics represent many rock types, akhough 
quartz, quartette, slate, schist, granite, pegmatite, gneiss, and porphyry pre- 

The beds dip vertically in the central parts of the Series "G", and at a steq> 
angle to the east iu both upper and lower parts. There are signs of compression 
throughout the scries, and some shearing stress h evident m parts. Cleavage 
planes are strongly developed, and these have a fairly constant dip of about 6j 
to the west throughout. 


The Western Outcrop ("Gl") 

The tillite "Gl" outcrops in the western fold o[ the anticline. It is found 
above the sub-glacial quartziles "I'T' in ;he central and western parts of the 
Campbell Range, and the northerly extension of this which runs through Mount 
Lock. No complete sections could be obtained from these areas, but examination 
of the basal beds of the scries revealed the same characteristics as those of U G", 
which have already been described. 

Isolated outcrops of the tillitc in its upper horizons showed that the angle 
of dip becomes progressively smaller to the west, until the series grades into the 
Tapley Hill series in the broad valley in which lies the township of Caltowie.. 
Structurally, this valley is a syncline, contrary to the neighbouring Jamestown 
valley, which is an eroded anticline. On the western margin of the Caltowie 
syncline the Tapley Hill series and tillitc once again outcrops in a ridge which is. 
the southern continuation of the Narrien Range. The latter contains ihe Appila 
Gorge, to which reference has already been made in discussing Series 'v' 
and "F*\ 

The Eastern Outcrop ("G2") 

The arkose grit in the upper part of the tillitc series "G M proved a useful 
indicator when working on 4, G2"> where it is a distinctive bed 45 feet thick. 
4 'G2" outcrops as the eastern fold of the syncline, in the Browne Hill Range — 
known locally as the "Cttnowie Melt." 

The tillite was traced to the north as far as Section 469, Hundred nf Wbyte. 
In this part of the range faulting and folding is very prevalent, and "there are 
numerous intrusions of quartz veins. Frequently associated with the quart/, is 
a siliceous iron ore. To the south, it was noted that the tillite continues for a 
number of miles south of the Hundred of Bclalie. 

The Jamestown-Terowic road follows a valley through the tillitc in the 
vicinity of Section 202, Hundred of Bclalie., This valley is the result of stream 
erosion in the zone of weakness following a small transverse fault. 

Tapley Hill Shales And Slates 

The beds composing this series are folded into a broad syncline with a north- 
south axis. This syncline extends from the tillite-contaming ridge *'G" east of 
Jamestown, to the tillite-containing Browne Hill Range *'G2", and has an 
approximate width of 5 miles. The syncline is responsible for the presence of 
a wide valley, which becomes broader and more clearly defined in a southerly 
direction towards Spalding. In the latter area the valley has a comparatively flat 
floor, hut proceeding north it is broken with increasing frequency by small rounded 
hills and ridges. These reach their maximum development in the North Bclalie 
district, where the valley floor is almost completely replaced by a series of 
alternating rises and depressions, The hilly parts are covered by a relatively 
shallow grey residual soil (Yangya silty loam) in contrast to the deeper red- 
hrown colluvial soils ('Belalie Scries) found in the depressions and on the valley 

On account of the persistent soil covering on the beds of this series, it was 
only after detailed investigation that a satisfactory number of outcrops showing 
dip's could be found. In general the rocks have well-developed cleavage planes, 
with an angle of 50° to K0° to the west, and these are frequently more clearly 
defined than the bedding planes. Towards the Browne Hill Range the cleavage 
and bedding planes are largely coincident, and this has produced some very fine 
rlajrs tones. 


It has already been noted in the discussion on the tillite series that the upper 
horizons of glacial beds gradually merge into beds of the Tapley PI ill series. This 
transitional zone is about 300 feet in thickness, and consists largely of fluvio- 
glacial slates in which occasional erratics occur. The lowest beds, which can be 
safely assigned to the Tapley Hill series, are thinly laminated argillaceous shales 
and slates, which in parts show ribbon banding. Most of these rocks are non- 
calcareous, but interbedded here and there are very thin bands of dark blue-grey 
impure limestone. These limestone bands appear to be similar to limestones which 
overlie the tillites in many other parts of the State, The basal shales and slates 
were mostly weathered to a soft yellowish-brown rock, while iu some cases slates 
were stained a deep purple. These basal beds of the series are similar in character 
on both sides of the syncline. 

The basal beds gradually merge into the more typical ribbon slates, which 
dominate the series. However, occasionally the banding is not clearly dcfined> 
and in some instances the slates are replaced by grey siliceous horn £ els. 

The banded slates are very fine-grained. They closely resemble the Tapley 
Hill slates in the Mount Lofty Ranges near Adelaide, and in the Flinders Kange 
further to the north. They are very similar to those described by Howchin 
(loc. cit.) at Orroroo, 

The true thickness of the Tapley Hill series in the localities examined is from 
10,000 to 11,000 feet. 

Banded Siliceous Limestones; 

In their upper horizons the rocks of the Tapley Hill series ("H") merge 
into impure banded siliceous limestones. Although outcropping in parts of the 
north Belalie district, they are best exposed about four miles east of Jamestown 
in Sections 319 and 320, Hundred of Belalie, where they are seen in a low hill 
rising from the main valley floor. 

The limestone has been used on a small scale for cement manufacture, and 
has been quarried in Section 319. The rock is Bne-grained, very hard and com- 
pact, and is dark blue-grey in colour. Banding is shown clearly on weathered 

A representative sample of the limestones was analysed, using the assump- 
tion that calcium and magnesium would be present as carbonates, and iron and 
aluminium as sesqui-oxides. The following results were obtained: 

Sio 2 

MgC0 3 - 

Auo 3 + PiyOj - 

Although the rock is characteristically siliceous, the percentage of silica 
shown here is much higher than would be given by limestones from specially 
selected horizons. The sample chosen for analysis was reasonably representative 
of the whole outcrop. A significant feature is the low proportion of magnesium 
compared to that in the analyses of limestones from series below the tillites. 

It is believed that these limestones, "J". Wtff represent the siliceous lime- 
stones below the Brighton limestones at both Adelaide and Orroroo, or else the 
basal beds of the Brighton limestones themselves. 





101 -3fc 



The folding which has produced the alternating anticlines and synclines is 
typical of a very large part of the Mount Lofty and Flinders Ranges. In this 
particular region, the fact that the pressure was not completely uniform is shown 
by the arcuate form which the outcropping beds now exhibit (cf. plan). 

There has been strike-faulting south-west of Jamestown, which has caused 
some repetition of beds in this sector. Repetition of part of the arkose sandstone 
and quartzite bed ("B") in the central part of the Hundred of Belalie has also 
been caused by strike-faulting. There is no evidence of strike-faulting along the 
lines of section ("PQ", "RS"). 

Although there has been a considerable amount of minor cross-faulting, there 
has been little on a major scale. The largest cross-fault has resulted in a displace- 
ment of nearly half a mile in the sub-glacial quartzites of the Campbell Range 
(cf. plan). Minor cross-faulting has been responsible for the development of 
zones of weakness in some of the beds, which has facilitated the down-cutting 
of streams. 


The writer is indebted to Professor J. A. Prescott for the opportunity to 
carry out the investigations, and to Sir Douglas Mawson for advice given from 
time to time during the course of the survey. 

Thanks are also due to Messrs. C. G. Stephens, R. Brewer and P. D. Hooper, 
of the Soils Division of the Council for Scientific and Industrial Research for 
their co-operation, and to Mr. L. M. W. Judell and Miss E. Gerny. 


1 Howchin, W. 1930 "The Geology of Orroroo and District." Trans. Roy. 

Soc. S. Aust., 54 

2 Mawson, D. 1941 "Middle Proterozoic Sediments in the Neighbourhood 

of Copley." Trans. Roy. Soc. S. Aust, 65 

3. Hossfeld, P. S. 1935 "The Geology of Part of the North Mount Lofty 

Ranges." Trans. Roy. Soc. S. Aust., 59 

4. Howchin, W. 1939 "The Geology of South Australia," 2nd ed. 

5. Jack, R. Lockhart 1919 "The Phosphate Deposits of South Australia " 

Dept. Mines Bull., No. 7 

6. Segnit, R. W. 1939 "Geological Survey of South Australia." Dept. Mines 

Bull., No. 18 

7. Sprigg, R. C. 1946 "Reconnaissance Geological Survey of Portion of 

Western Escarpment of the Mount Lofty Ranges." Trans. Roy. Soc. 
S. Aust., 70 



By Reg. C. Sprigg 


During the course of the second World War, officers of the Hydrographic Survey Branch of the 
Royal Australian Navy discovered several huge gashes along the normal course of the continental 
platform of New Guinea in the Morobe area. These are submarine canyons. More recently, 
following an official request to the Navy on the writer's behalf, further examples have been locatd 
off the South Australian coast (fig. 1). 



By Reg. C. Sprigg * 

[Read 9 October 1947] 






General Features of Submarine Canyons 

Theories to account for Submarine Canyon Formation 

Low Sea Levels of the Pleistocene and possible Canyon Relationships . . 
The New Guinea Submarine Canyons 

Coral Reef Relationships and Sea Level Variations 

The Murray Submarine Canyons 

Continental Platform in Relationship to the Submarine Canyons 

Reconstructed Possible History of the Murray Canyons 



During the course of the second World War, officers of the Hydrographic 
Survey Branch of the Royal Australian Navy discovered several huge gashes 
along the normal course of the continental platform of New Guinea in the Morobe 
area. These are submarine canyons. More recently, following an official request 
to the Navy on the writer's behalf, further examples have been located off the 
South Australian coast (fig. 1). 

The New Guinea sub- 
marine valleys have been 
(named respectively (from 
west to east), the Waria, 
Eia and Gira Canyons, 
after the rivers which enter 
the sea near their heads. 
The Murray examples are 
referred to as the west, 
central and east submarine 
canyons, and they occur 
south of the centre of 
Kangaroo Island. Their 
association with the River 
Murray at the time of de- 
velopment is inferred, but 
the case is not yet proved. 
It is hoped that more light 
will be thrown on this pro- 
blem when several more 
echo - sounding traverses 
are completed in their 
vicinity later this year. 




G.PK del. 


Fig. 1 Locality Plan 

* Assistant Government Geologist, South Australian Geological Survey. 
Trans. Roy. Soc. S. Aust., 71, (2), 1 December, 1947 


It is emphasised that, whereas the New Guinea examples have been covered 
extensively by sounding- traverses aggregating several hundred miles in the 
immediate canyon vicinity, the South Australian examples are known only from 
u single traverse. This was designed by the writer to locate the 100 fathom line 
more accurately and to seek possible canyon structures which could be related to 
the River Murray. 

More than 100 submarine canyons are known from various parts of the 
world, ami the list is steadily increasmg. These structures have many features 
in common, viz.:- — 

1. Distribution is world wide. 

2. The furrowing is obviously the work of running water. 

3. Most of the erosion appears to have been in soft watcrsoaked sediments. 

4. Many submarine canyons are obviously genetically related to adjacent 

5. Their greatest development occurs at the "fall of!" from the continental 

6. By analogy wirh suhaerial valleys, they are youthful features of the earth's 

7. (jencral similarities of canyons surest simultaneous formation. 

8. They are definitely post-Pitoccnc features, as^ evidenced by youngest, wall 

Numerous theories have been put forward in an endeavour to account for 
these interesting topographic phenomena, One school considers them to have been 
subaerially eroded, while another regards them as a result of submarine currents. 
In this paper the main theories will be outlined very briefly, and particular 
attention will be paid to that, theory concerned with sea bottom density currents, 
which is the one favoured by the writer. In malting this latter statement, it is 
recognised that under special circumstances various processes may have inter- 
acted in canyon erosion. For example, vv'hcre the canyon walls arc steep sub- 
marine slumping of sediments would almost certainly take place, and these effects 
could be imensified by seismic activity, the net eilect being increased erosion. 

(a) Regional Uplift of the Land 

This theory postulates world-wide continental uplift at the time of canyon 
formation to extents, of 5.000 to 10,000 feet. This would expose the continental 
*Iope to powerful suhaerial stream erosion. Such a theory has little, if any, 
factual basis. The earth's crust is aJmost certainly too stable to allow of such 
rapid and extensive land movement. Even supposing such movements could 
occur, it is considered very unlikely ihal the return to the present state could be 
accomplished without much relative warping- along the strike of the continental 
platform, producing much greater irregularities of deptii at the outer continental 
shelf margin than is apparent today. 

(b) Temporary Kustatic Fall of Sea Level 

A geologically recent, world-wide lowering of the sea level of up to 10,000 
feet, would perhaps account for the formation of the structures by suhaerial 
erosion. However, the most extreme temporary sinking of ocean basin floors, 
superimposed on effects of glacial eustasy, would have "to he involved, and, np 
to the present, there is no reliable evidence for such j*rc.-ir vertical movements 



within recent times. Eustatic lowering of the sea level (Zeuner 1946) is 
estimated to account for a fall of only about 55 fathoms (330 feet or 100 metres) 
at the height of Pleistocene glaciation (by extracting water from the sea and 
(vmg it up m ice caps). Such a lowering of sea level will obviously explain the 
existence of many river channels across the shallow portions of the continental 

(c) Currents of the Ocean Bottom 

There are several theories based on the 3bove. Some are very Ingenious, 
but thev appear to be unsupported by fact. For example, Douglas Johnson 
(1939), suggested a special type of current called "spring sapping/ 7 which 
assumes that water from subaerial sources, under special circumstance, seeps into 
submarine (continental platform) sediments and escapes somewhere on the 
continental slope. The water is thought to issue with dissolving power and kinetic 
energy sufficient to erode canyons. Another theory by W. H. Bucher, 1940, 
postulates reflex currents generated by powerful earthquake waves in the ocean. 
However, as many of the largest canyons occur in areas least effected by earth- 
quake shocks, this theory is of little importance. 

The most satisfactory theory, at least to the author's mind, and one which 
is very strongly supported by field analogy and laboratory research, is one by 
K. A. Daly (1942), which is based on special types of bottom density currents in 
aqueous environments. The canyoning is considered to be brought about hy 
gravitational muddy bottom currents temporarily endowed with a density greater 
than that of the normal water of the overlying medium. Daly suggests further 
that such muddy bottom currents were a feature of the Plehtncene glacial period*? 
when the sea margin was closer to the fall off from the continental shelf, and 
when much of the earth's climate was decidedly more pluvial in character than 

.Studies in natural and artificial lakes (e.f/ t , Lake Geneva in Switzerland and 
Lake Mead Reservoir in U.S.A.) indicate that muddy suspensions of the type 
mentioned are by no means uncommon,, and that their powers of erosion are 
considerable. They are capable of deep excavation, and they frequently move 
quite coarse gravels along the sloping lake bottoms. 

Dr. P. H. Keunen, of Holland (1938). has shown by laboratory experiment 
that silt-laden bottom currents can erode and maintain themselves and become 
self -accelerating down slopes typical of the continental shelf. Keimeti conducted 
his experiments in a large tank with bottom topography designed to represent a 
typical section across the continental shelf and continental slope. lie found, on 
releasing muddy suspensions on the upper shelf area, thuL they gravitated down 
the continental slope, and that if a slight crease existed down such a slope, silt- 
laden water was pirated from cither side, thickening the current along the crease 
and increasing its velocity. Moreover, if the bottom was covered by clay, the 
density current became turbulent beyond rhe continental shelf "fall off/' causing 
the current to erode deeply and thereby adding to the current's density, mass and 

Knowing the slope factor and the density of the silly suspension, and by 
measuring the velocity and the so-called hydraulic mean depth of the current, 
Keunen arrived at an empirical formula which can be used to calculate the velocity 
of given suspensions down particular continental slopes. 

Applying this formula, Daly has found that on a continental slope of 1 in 15, 
and w'th an elective density of the flowing water and silt mixture of 0*0005, 
the velocity is about 4 miles per hour. This would be sufficient to sweep along 


coarse gravel and, therefore, to erode silt and sand with great case. In this way, 
there is an effective means of submarine erosion available for canyon excavatiun 
under the right conditions. 


According to the theory of glacial eustasv, the low sea levels of the Pleisto- 
cene were due to the extraction of water from the sea caused by accumulation of 
ice, particularly on sub-polar landmasses of the Northern Hemisphere. According 
to Zcuner (1946), and other authorities, the last glaciation reduced general sea 
level by about 100 metres below its present stand (the Frc-Plandrian Regression). 
The geographic consequences of these various changes in sea level, correlated 
with at least four glacial periods, must have been far-reaching. Coastliue con- 
figurations must have changed greatly and rapidly, while rivers were at the same 
time alternatively hetrunked or engrafted- In addition, coastal features were at 
the same time drowned or stranded. 

In the present case, our concern is with ihz lower stands of .sea level, which 
resulted in coastal advance over considerable widths of the continental platforms 
uf the world, Muddy river waters were now carried far across the continental 
platform and closer to the steeper slopes of the continental shelf before encounter- 
ing the sea. At the same time, the related change of climate and relative alti- 
tudinal increases of the local landmasses would tend towards increased pluviality, 
at the same time swelling the volume of local rivers and consequently increasing 
their sedimentary load. In most situations the soft sediments of the partially 
exposed continental platform supplied further material for river transport, both 
by traction and in suspension. Ocean waves also had access to these sediments, 
extracting much material in suspension, The scene seems now to have been set 
for the operation of the most active canyon-forming processes. 

The various muddy suspensions, and particularly those associated with 
rejuvenated rivers, provided the bottom density currents, which, according to Daly 
and others, in many cases were able to gravitate and maintain themselves across 
the narrowed shelf zone to the greater slope beyond the shelf edge. This, in turn. 
accelerated the suspension currents causing local canyon formation. Tn this w r rty 
many furrows were produced on the shelf edge, but, particularly where muddy 
rivers entered the sea, deeper canyon structures were formed. 

The New Guinea submarine canyons are quite typical examples of submarine 
valleys which can be related very satisfactorily to local river mouths (fig, 2). 
They reach the large dimensions for which such .slrnrhn^s are noted, and their 
courses lie fairly directly across the continental platforms. 

They differ somewhat from the celebrated eastern United States examples 
iti that their respective canyon heads approach within less than one mile or so 
of the modern coast. In their close relatonship with river months, they arc akin 
to the Congo type of submarine canyon (sec Veatch and Smith, 1939). 

Statistical details of the canyon structures arc summarised in Table I. It is 
seen that lengths of the various canyons from their "heads" to the (projected) 
continental shelf edge vary from 7% to 10 miles and that separation nf the "heads" 
from land is between O-S and 12 miles. Thi.i information, considered in die. 
light of the local widih of the continental platform (7 to 9 miles), indicates the 
relatively direct courses taken by the canyon beds. At the shelf edge the struc- 
tures range w width between 2 and 4 miles and their Tcspectfve beds lie from 


















I v* 




! > 



1 ' f* 





! < 






! X 


a- t 








I < 

g o 





f> <_< i 



4,500 to 4,800 feet below sea level in the same region (fig. 3). The canyon 
bottom gradients are relatively steep, averaging between 1 in 10 and 1 in 13 
(i.e., 10-0 and I'l^o). Little' is known of canyon development beyond the con- 
tinental shelf edge, hut the structures obviously continue strongly down the con- 
tinental slope to abyssmal depths. 

Fir. 3 

Section through submarine canyons of thr Morobe urea, drawn along the 

projected continental shelf edge. The portions of the section remaining above 

50 fathoms are barrier (coral) reefs. 

Available information indicates that the sea bed and canyon walls generally 
are of soft volcanic mud, except in shoals where it is coral. The muddy sediments 
are excellent media for swift erosion and have certainly aided the rapid enlarge- 
ment of the canyons, laterally and longitudinally. 

The Kia submarine canyon differs from the other two in that its present 
head does not oppose the modern mouth of its parent river. The old river outlet 
is still to be observed opposite the canyon, but the new mouth is further west- 
ward and is not opposed by a secondary canyon. 

In view of the close relationship with rivers entering the sea locally, statistics 
of the respective parent river development, namely trunk stream length and basin 
area, have been included in Table L As the area is one of high rainfall (Edie 
Creek and Kokoda average 107 and 143 inches per annum respectively), it is 
obvious that huge volumes of water are discharged into the local seas annually. 
The highest average monthly rainfall exceeds 16 inches and probably during the 
Pleistocene glacial periods was much greater, giving some indication of possible 
erosive or transporting power of the local streams under most favourable condi- 
tions. Turbulent waters such as these, crossing an exposed sea bed of volcanic 
ash. would certainty become extremely well loaded with sediment, sufficient to 
form powerful density currents. 

Tmh.e T 
Submarine Canyon Statistics (New Guinea Group) 


Ek - 














Width at 

edge of 


shall' edge 



3 to 4 










U 9*6 

1 : 12-7 

of 100 
line to 
land at 
canyon head 




Modern "'Paruut" 


Area of 








Coral Reef RflkArjnNsmKs and Sea Level Variations 

^Tnvsive coral reefs skirt the outer margin of die continental plalform in die 
Morobe area. Between (hem and the shore line, small isolated coral colonies are 
common, The outer barrier is breached h v the submarine canyons which arc 
lined on either side by irregular but extensive coral reefs. 

The outer reef base extends to at least 70 fathoms below present sea level, 
and as deep water lies between these corals and the land, they appear to be 
typical barrier reefs. Nothing is known of their present condition of growth, 
ie. r whether their general sunken aspect is due to recent negative land or shelf 
movement, or whether the corals are dead and cannot therefore build up to the 
modem sea level. 

According to Vaughnn and Wells (2943) and Yonge (1940) reef corals will 
not flourish in depths exceeding 2S-30 fathoms, the controlling facior here heinR 
light penetration. Under these controlling circumstances, reef coral* could not 
therefore have established themselves on the outer margins of the coutiTicuml 
shelf under the modern high sea level. As the shelf adjacent the reefs exceeds, 
on occasions, 70 fathoms, a lowering of the sea surface by 40 to 50 fathoms may 
be assumed for their establishment (unless equivalent land movements could be 
proved). This assumption is in keeping with figures for the Pleistocene prc- 
Flandrian regression,* * > and ii therefore seems probable that the coral rect 
development commenced at this lime or during an even earlier Pleistocene 
(glacial) low sea level. 

Furthermore, on the assumption that the formation of all submarine canyons 
was contemporaneous (post- Pliocene), it can be inferred that harrier reef forma- 
tion could not have prc-dated this period a? such reefs would have dammed back 
and destroyed bottom density currents, and therefore prevented canyon excava- 
tion. This is in keeping with Gardiner's theory (1931, p. 115) that l, there arc no 
indications of any true reefs living or elevated before late Tertiary (Ccnozoic) 
time." According to Daly's theory, discussed previously in this paper, the lower- 
ing of sea level (here considered necessary for the introduction of coral growths 
at the shelf edge) would also have been the most favourable for submarine 
canyumng, due to increased pluviality and to exposure of soft continental plal- 
form sediments, Jt seems reasonable, therefore, to assume that the formation 
of submarine canyons and the establishment of barrier reefs was contemporaneous. 

Another condition for thriving coral growth is that (lie local water should 
not be muddy, as muddy conditions reduce light penetration, and smother fjjij 
coral animal. If it is tu be assumed that submarine canyon and reef establish- 
ment occurred during the low sea level phases of the Pleistocene, it would appear 
that muddy suspension currents from coastal waves as distinct from river action 
on exposed continental platform sediments, were insignificant in this area, at 
least for portion of the period of low sea level.'^ If the former muddy currents 
had been widely developed during the shallow sea level stand, the shelf edge reefs 

< l > The tail of sea level, which was contemporaneous with the Last Glaciatioit, 
probably attaining almost 100 metres (approximately 330 feet or 55 fathoms) below 
modern sea level. 

W Daly (1935) has suggested that during the earlier periods of the respective 
Pleistocene low sea levels, the muddy fractions of the outer continental shelf sediment* 
were winnowed out to be deposited in deeper, quieter watery leaving less muddv sedi- 
ments in the littoral zone. This is logical, but it would suggest that cation furrowing, 
if resulting purely from bottom density currents tnitiatcit by shore wave action, would 
be active only for short periods during the successive low sea levels. This surely would 
reduce their significance considerably. 


could net have been established due to the muddiness of ihe bottom waters, even 
though other -factors may have been favourable. It suggests, 1 here t Are, tliat the 
density currents responsible for most of the canyon excavation were initiated by 
silt-laden river waters entering the sea in confined zones. According to Keurten 
(see earlier), once significant channels are formed down submarine slopes, density 
currents become moie restricted, and in this way adjacent stretches of sheli edge 
would remain free of silty suspension cumins, thus allowing establishment of coral 
growths while depth and conditions, were satisfactory, At the same time, it is 
obvious that coral growth within the course of the muddy density currents would 
be completely inhibited. Hence the presence of gaps in the barrier reefs at the 
point where the currents discharge across the outer edge of the continental shelf. 
These gaps would be subsequently widened by undermining and collapse in the 
normal course o£ canyon development, The two processes would also affect reefs 
skirting the margins of the canyons, but here there would he a tendency to 
restriction of Lateral expansion and a steepening of the canyon walls. 

It has been suggested previously that the low sea level stands, necessary for 
the establishment of coral reefs in such positions* were correlated with one or 
more Pleistocene glaciatious, and, as reef corals do not flourish in waters below 
20° Centigrade, a minimum temperature of this order seems probable, However, 
without detailed knowledge of reef sections, which may reveal horizons of 
repressed or completely inhibited coral growth, this can only be conjectured. 
Coral growth may have been destroyed locally, to be re-established only when 
the seas warmed again during subsequent inter glacials As the minimum average 
monthly temperature of the local coral seas, appears to be at least 25° Centigrade, 
such a marked lowering of temperature is improbable. 

From Ihe foregoing discussion, it Is obvious that the Antecedent Platform 
theory of the origin of coral reefs (Hofrmdster and Ladd, 1944), is applicable 
tn this area if it is correct to infer that Pleistocene* sea levels fell by as much as 
40 to 50 fathoms. 'This theory states that any bench or bank that is located nt a 
proper depth within the circum-equatorial coral reef zone is potentially a coral reef 

Tn considering the survival of coral reefs m competition with the rising sea 
level at the onset of a severe interglaoal- it is to be noted that Daly (1935) has 
calculated that the rate of rise, probably never exceeded 3 mm. per annum. This 
figure is well within the range of reef upgrowth, Gardiner (1903) concluded 
from his experiments that normal upgrowth may be 27 to 45 metres per thousand 
years, and later studies have confirmed such phenomenal growth rates- As a 
corollary of this it would appear that modern coral reefs evolved to their present 
massive proportions only with the abnormally large (and relatively rapid) 
fluclnations of the rieistoccne sea level, Such fluctuations would alternatively 
reduce pre-existing reefs with falling sea level, and initiate new ones with the 
rising. The rapid sea level upgrade would facilitate maintenance of the prodigious 
food supply necessary for rapid and healthy reef upgrowth. 

Speaking generally, there is no positive evidence from any part of the world 
that ne?v submarine canyons arc being formed under the present high sen level 
stand, but it is quite possible that some well-established canyons arc still being 
extended, especially in areas where the canyon heads approach close to the shore 
and pluviality is relatively high — as in the Moro'be area. In this case, two' of the 
canyon heads <Gira and Waria) approach within less than a mile of their respec- 
tive river mouths (although the local continental shelf is about 10 miles wide), 
thus providing very steep sea-bottom gradients tor silly suspension currents. In 
the thin! instance, that of the F-ia submarine canyon, the canyon head does not 
oppose the mouth of the modern Eia River, although an older oudct of the same 


river obviously once did so. There is no evidence that the modern outlet is 
producing a new submarine valley, even though the 100 fathom line lies relatively 
close to the river mouth. This may suggest that either the shelf slope from the 
newer mouth, although relatively steep, is still insufficient to maintain bottom 
suspension currents over the distance, or that tlte amount of material in suspen- 
sion, produced by Eia Rivet waters, is row insufficient to produce significant 
density currents. There is the further possibility that well-established coral shoals 
which effectively block and destroy any gravity currents intervene at the edge 
of the local 100 fathom line, It may be that ail three factors are operating 

In the foregoing discussions little has been said of possible large vertical 
movement of !he New Guinea land mass intensifying or diminishing Pleistocene 
variations in sea level. The fact that the outer continental platform is relatively 
constant at 60-70 fathoms below sea level suggests that the region has not moved 
vertically very much in relatively recent times. The outer depth figure shows 
fair correspondence with similar continental platform relationship elsewhere in 
the world, particularly in the coral seas. Nevertheless, this aspect is one which 
demands close field investigations, as there arc reports of late Tertiary or Pleisto- 
cene coral reefs now raised many hundreds of feet above sea level in parts Of 
New Guinea (see David 1932). 

It is noticed that the continental platform is generally shallower to the south- 
cash and this may argue in favour of at least minor local warping movements in 
relatively recent times. This aspect may repay further investigation. 


The present discoveries are the first to be found along the Australian coast. 
Original soundings in the canyon vicinity by the Admiralty were obviously very 
scattered, and even if odd very extreme soundings had been recorder! at the 
canyon site, they would almost certainly have been disregarded ifl the preparation 
of sea-charts on the grounds that they might have been inaccurately located. 

The little information at present available on the submarine canyons i> limited 
to a single "continuous" echo-sounding traverse across them (fig. 4). The 
traverse was designed to follow the 100 fathom line, as this was considered to 
offer the best means of locating possible canyons It ran from opposite the 
western end of Kangaroo Island lo near Bcachport, a distance of almost 180 miles. 
As a result of the survey, the platform edge has been found to be remarkably 
regular, broken by major sliucturos in only one locaKly. approximately due sotrth 
of the centre of Kangaroo Island. There is little or no doubt thar these structures 
aic submarine canyons, as the wall slopes reach at least I in 4*7 as against the 
continental slope hereabouts of 1 in 20 or less. 

It is not yet proved that the canyons are genetically related to the River 
Murray, but there is good reason to assume such a relationship. The presence of 
three canyons closely spaced on the shelf edge, which otherwise is remarkably 
even, suggests that one process or one group of processes only has controlled 
canyon excavation, rather than that the process has been a fortuitous one. The 
mo?t likely control seems to have heen. by an ancient river entering the sea locally 
during a low sea level stand. The only large river available would be the River 
Murray, although, of course, a secondary river from the drowned St. Vincent 
Gulf bed could have entered the sea in this region, but it seems more likely that 
such a river would be engrafted on the extended River Murray. 

The three separate submarine valleys occur in a shelf edge distance of only 
three miles. They range to at least 750 fathoms (4,500 feet) at the survey Vint, 
and almost certainly continue down the continental slope to abyssal depths 



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Murray Submarine G\nion§-%5 


11 — £ 

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Plan 4 

Plan o{ the continental ihclf, showing major physiographic features m the 
auUaianne canyon vicinity. Sites *'A" and "B" on the fathometer traverse 
from Robe to Cape WOloughbv arc Ch$ low poiuts indicated in fig. 7. 

extent landward is completely unknown, but it is unlikely to be extensive. The 
(Jeffrey Deep which lies to the south exceeds 3,000 fathoms). The canyons" 
*cparate submarine valleys are so- close together that definite ridges occur 
between two of them (rig. 5), From west to east, along the tme of traverse, they 
range in depth to 550, 625, and 750 fathom 3 respectively. 

The Continental Sttllf and Can vox Relationships 

The continental platform is relatively narrow in the Beachport-Robc coastal 
run, Tt varies in width from about 16 miles opposite Cape Banks to 30 or more 
opposite Cape Jaffa. From this latter point the shelf widens to more than 100 
miles, due to the westerly sweep of the 100 fathom line and to the marked coastal 
concavity associated with the Coorong, The shelf narrows again very consider- 
ably west of Kangaroo Island. A sketch section across the continental platform 



.... ... m 

S£A L£\/CL 

-200 jf 

1 A • 








(y* Z£ \l 





f f 1 



C A L E 

Fig. 5 
Profile section ("A") through the Murray Canyons. Note steep wall 
gradients as compared with the general gradient of the local continental 

slope ("B"). 

at its widest development is given in fig. 6. The very approximate grades of the 
critical sections of this shelf are as follows: — continental shelf, 1 in 880; and 
steeper continental slope 1 in 20. 








— Jeffrey Deep 


Con f/"/7 errtft Sfte/f 

Con///7e/7f&/ Sfopc 
Figures indicate depth in fathoms 

Fig. 6 

Generalised section across the continental platform fronting Jeffrey Deep. 

The true shelf, as indicated, is more than 100 miles wide and represents 

conditions opposite the Murray River mouth. 

Fig, 4 has been produced from admittedly insufficient data. The soundings 
used in the interpretation of the form line are mainly from Admiralty charts, and 
while these are distributed more copiously nearer the coast line, they are far too 
few in deeper waters. Hence, while the plan probably gives a reasonable general 
picture of shelf development, il must be considered as unreliable in detail. The 
representation then is partly conjectural, but it is felt that it is a reasonable 
approximation of the natural features. It is hoped that in the coming season 
several more traverses will be made across the zone, and that a more definite 
picture of the structures will thus become available. 

The lines of the two traverses already made across the continental platform 
are included on the plan, and the sea floor detail from the Admiralty charts has 
been modified slightly in accordance with these. The first to be run was the 
Robe-Backstairs Passage traverse, but it failed to locate any definite "gorge" in 
a favourable situation for a possible drowned River Murray course. However, 
at least two possibilities apparent on the traverse (fig. 7) are indicated, and 
through these two points alternate courses for the inferred drowned section of. 
the River Murray are drawn on tig. 4. Minor seaward concavities in the 
40 fathoms form line have been used in designing these two courses, but it is 
realised that these form line deviations themselves represent only the best one 
can do with such a paucity of soundings in this area. The immediate subcanyon 
area is not covered by Admiralty charts, so the amount of interpolation in this 
area is obvious. 

fATHQMS £ ea 



Loup Soundi ng Trm'lpse. 



Suggest? d AffpsnzTh* L&c a t&ns A i-S, 
iQC^SflSS HfW River. 

30 MILE.5 *0 


Fig. 7 

The great exaggeration of the vertical scale in relation to horizontal cli&t&nce 
is to be noted carefully. The ,sea floor over much of the section is extremely 
flat; it varies less than 30 feet vertically from the 30 mile To (h* 80 mile 

indications on the section. 

Shallows occur immediately north of the submarine amy on s, and these may 
be associated in some way with Pre-FIandrian shore-lines. They do not come 
within less than 5 fathoms of sea level (South West Rock is the shallowest) and 


tbc nature of the rock is unknown. It is possible that it may be consolidated 
beach sand-dune material, such as that which occurs at Wedge Island at the toot 
ol Spencer Gulf. If this is proved, it would be additional evidence favouring an 
old coastline stand at about 40-50 fathoms ( ? Pre-Flandrian Regression). 

Form line* in St. Vincent Cult area indicate that there is a fairly definite 
longitudinal depression line in tins region. It is most probable that there was a 
rivet in this 7.oue during the Pleistocene low sea level stands, and that this river 
Unwed south, pos>ibly to join the River Murray near Sanders Rank. A sounding 
traverse has been run from Port Adelaide to Troubridge Lighthouse in an 
endeuvout to locate an actual river course, but without success However, con- 
siderations of gul* bottom topography will be discussed in a later paper. 

In the continental platform area facing the Southern Ocean, it must always 
be borne in mind that littoral wave action and other blanketing processes may 
have effectively destroyed much or most of any old submerged river courses. 

Concerning the inferred submarine canyon •'detail," the valleys have been 
represented diagrammaticatly as parallel structures trending down the continental 
slope. This appears to be the safest assumption in view of the information avail- 
able, It is realised though that there is- every possibility that two or all the struc- 
tures may unite at sonic lower point or points on the continental slope, in den- 
dritic fashion. 

Possible Historical Development of Tin: Ml'krav Canyons 

Thrilassostalic terraces (see Zeuner. p. 130) at: Murray Bridge indicate that 
within relatively recent times die l.cfc&l base level (and therefore sea level t has 
bcen at least 100 feet below its present position. The river bed in this locality may 
have been eroded even lower if granite bars -had not intervened preventing rapid 
deepening in that locality. 

This lowering of base level is in keeping with the postulated low sea level 
stands, of the Pleistocene epoch which, according to Zeuner (1946) were as low 
as 100 metres (330 feet or 55 fathoms) below present sea level datum. With 
the retreat of the sea level to this extent the coast migrated seaward (Pre- 
Flandrian coastline, see fig. 4) causing the River Murray, and possibly other 
streams to advance over soft continental shelf sediments. The most direct course 
from the modern Murray mouth region to the sea would bring the Pre-Flar.drian 
river mouth to a position about 40-50 miles east of the Murray Canyons, It 
:cems probable, therefore, that if the canyons were directly related to an old River 
Murray mouth or mouths, the river naist have pursued a much more westerly 
course'than that of the shortest route. As gradients of the local continental shelf 
average I in S&0, and in many places are far lower, it is not surprising if the 
course did wander considerably. The river apparently hugged the Mount Lofty- 
Kangaroo Island horst structure to some extent, much in the manner that the 
existing river hugs the Mount Lofty horst today. 

Such a course may have been fortuitous or controlled to a degree by minor 
■nurth downward movements, known to have occurred in south-east^ sub-coastal 
regions during the Pleistocene. Littoral drift may also have operated, tending to 
deflect the river mouth north and west with the retreating sea. However, the 
case for north-westward littoral drift along the south-east coast, at least to the 
author's mind, is far from proved, and will not be emphasised here. 

A feature of the Murray canyons is their triplieity. This is particularly 
noticeable in the absence of other large submarine canyons either to the east or 
west along the shelf edge surveyed. They are unique in that only very small 


distances separate each of the three canyons. From the western boundary of 
the most westerly canyon to the eastern edge of the most easterly is less than three 

There are at least two distinct sets of conditions which could have led to 
such development. In the first place, three separate contemporaneous outlets can 
he postulated, related either to three separate Rjffge rivers, or to three major dis- 
tributaries of a delta system of the ancient (Prc-Flandrian ) River Murray. 
However, it is difficult to imagine three separate and contemporaneous large rivers 
entering; such a short stretch of coast, when for a distance of at least one hundred 
miles to the east there is no further evidence (?.#-, submarine canyons) nf such 
outlets. Nor does it seem prohahle that in delta formation three distributaries 
would be maintained for a sufficient time to establish submarine canyons. If 
such a delta formation is assumed to have existed, the probable effect would not 
have been three separate canyons* bul subparalkl suhmarine furrowing with a 
tendency towards pirating by a major furrow and the ultimate formation of a 
single large submarine canyon. 

The second hypothesis is that the canyons were formed in association with 
three complete changes in the course of the Murray, correlated with particular 
low sea level phases of the Pleistocene. It can he assumed that later canyons 
were formed by outlets completely separate from the "sphere of influence" of the 
tildcr canyon (s), as any course which discharged its load within the vicinity nf 
an established submarine furrow would have that load pirated by it, 

A further problem is the relative age of the submarine valleys. This could 
possibly' be determined only by detailed core sampling of the canyon walls for 
Vossil data. In is interesting to note that the easternmost canyon is the. deepest, 
and that the two to the west are progressively shallower along the line of section. 
'this suggests that the cast canyon has been subject to more prolonged erosion 
by bottom density currents. The later currents would have been products of 
cnastai wave action during subsequent low sea level stand, although the possi- 
bility of enlargement by still a further low sea level and river association cannot 
be overlooked. 


Six new suhmarine canyons are recorded. Three of them occur in the Morobe 
area of New Guinea in close relation to modern or old river outlets. They are 
eroded in volcanic muds (and probably also marine Tertiaries) across a conti- 
nental shelf only 10 miles wide, which descends to 50 and 70 fathoms at its outer 
limit. The continental shelf vdgc and the canyons themselves are lined by coral 
growths, and from this, on the assumption that reef corals cannot flourish in 
water deeper than about 25-30 fathoms, it is inferred that sea level had fallen 
probably 40-50 fathoms at the time of cairyon formation, sufficient to allow 
barrier reef establishment ai the shelf edge. However, in these estimates no 
allowance is made for possible land movements, These submarine canyons arc 
of typically grand dimensions and relevant data concerning their development is 

The Murray submarine cauyons are the first to be discovered bordering the 
Australian continent. They are considered to be related to the modern River 
.Murray, although the evidence for this is not complete. Three separate canyons 
are apparent in a width of less than four miles. Two alternative theories are 
advanced for canyon triplicity, both having relation to the repeated sea level falls 
of the glacial phases of the Pleistocene. 



The writer wishes to thank the Department of the Navy for supplying 
Fair Charts of the Morobe area and for undertaking several survey traverses in 
strategic positions in South Australian waters. Also, indebtedness is expressed 
to Lieut.-Commander Little of H.M.A.S. "Lachlan" for the keen interest dis- 
played in the search for submarine canyons, and to Dr. L. K. Ward for helpful 
suggestions during the final stages of the preparation of the manuscript. 

Bucher, W. H. 1940 Bull. Geol. Soc. Amer., 51, 489 

Daly, R. A. 1935 "The Changing World of the Ice *-\ge." Yale Univ. Press, 

New Haven 
Daly, R. A. 1942 "The Floor of the Ocean." Univ. of North Carolina Press 
David, T. W. E. 1932 "Explanatory Notes to accompany a new Geological Map 

of the Commonwealth of Australia." Sydney 

Gardiner, J. S. 1903 "The Fauna and Geography of the Maldive and Laccadive 
Archipelago." Amer. Journ. Sci., 16 

Hoffmeister, J, E., and Ladd, H. 3* 1944 "The Antecedent-Platform Theory." 

Journ. Geol., 52, (6) 
Johnson, D. W.. 1939 ''The Origin of Submarine Canyons," New York 
K2UNEX, P. H. 1938 Geol. Mag., 75 
Vatjghan, T. W., and Wells, J. W. 1943 "Revision of the Suborders, 

Families and Genera of the Scleractinia." Geol. Soc. Amer. Spec. Pap., 

No. 14 

Veatch, A. C, and Smith, P. A. 1939 "Atlantic Submarine Valleys of the 
United States, and the Congo Submarine Valley." Geol. Soc. Amer. 
Spec, Pap. ; No. 7 

Yonge, C. M. 1940 "The Biology of Reef Building Corals." Brit. Mus. (Nat. 
Hist.) Grt. Barrier Reef Exped. Rept., 1, No. 13 

Zeuner, F. E. 1946 "Dating the Past/' Methuen, London 



ByH. B. S. Womersleyand I. L. Ophel 


During a post-sessional excursion following the 1947 Australasian Science Congress in Perth, a 
remarkable number of the Characeae was collected by the first author from a small, shallow swamp 
on top of the peneplain of the "break-away" country above the Irwin River, near Minginew, 
Western Australia. The habit of the plant, and the enormous size of the cells, was unlike any of the 
Characeae commonly found in such localities. Most striking, however, was the complete absence of 
stipulodes, bract-cells and bracteoles, a characteristic of no previously described genus of 



By H, B. 5. Womerslk/ and I. L. Orziti,* 
[Read 13 November 1947] 

During a post-sessional excursion following the 1947 Australasian Science 
Congress in Perth, a remarkable number of the Characeae was collected by the 
first author from a small, shallow swamp on top of the peneplain of the "break- 
away" country above the Irwin River, near Minginew, Western Australia. The 
habit of the "plant, and the enormous size of the cells, was unlike any of the 
Characeae commonly found in such localities. Most striking, however, was the 
complete absence of stipulodes, bract-cells and bracteoles, a characteristic of no 
previously described genus of Characeae. 

Nandor Filarski (1937) described (in a Hungarian journal) two Western 
Australian Charophytes from specimens sent to him by G. Q. Allen. These 
specimens were from the collection of J. Groves at the British Museum,, and had 
been collected originally by Miss N. T. Burbidge. Filarski founded a new genus. 
Charina, on one of these specimens, although he had no fertile material (Sec 
later notes.) The other specimen he referred to Nitellopfis as N. inflata Fitarski 
and Allen . Unfortunately, it has not been possible bo examine material of 
N. inflata, as no specimens exist in Australian herbaria, and Filarskt's figures 
(reproduced in fig. 2) are inadequate in many details. From Filarskfs figures, 
however, N, inflata appears to be closely Tclatcd to our own species, also showing 
complete lack of stipulodes. bract-cells and bracteoles, but differing in several 
important details (see later). 

It is therefore proposed to find a new genus, Protochara, with P. australis 
n. sp. as the type, and to transfer rV, inflata to this genus as P. inflata (Fil. and 
Allen) comb, nov* 

Protochara australis it. sp. 

Plant dioecious, totally ecorticate, to 10 cm. high, light green in colour, with 
no calcareous incrustation; attached by branched, colourless, several-celled 
rhizoids (fig. 1 B). Stem stout, simple or with a few axillary branches, bearing 
4 to 7 whorls of branchlets; stem internodes 1 to 3 cm. long, 0*9-1*5 mm. thick. 
Branchlets stout, slightly incurved, in whorls of 4 to 7, of 3 or 4 segments; 
terminal segment consisting of a small mucronatc cell, 110-200/* long, ringed at 
the base by 5 or 6 peripheral nodal cells (fig. 1 D) ; subterminal segment large, 
2-3 mm. long, 1-1| mm. broad, asymmetrically inflated with the inflated side 
distant from the stem (fig. 1 A, B, D) ; intermediate segments -£-1 cm, long. 
almost as thick as the stem, slightly inflated when young. Nodes consisting of 
10 to 12 peripheral cells surrounding a plate of inner cells (fig. 1 C, D, G). 
Stipulodes, bract-cells and bracteoles completely absent Chloroplasts minute, 
forming vertical series in the cells (fig. 1 C). 

Oogonia verticillate in the axils of the upper whorls of branchlets, or borne 
singly or geminately at the nodes of upper branchlets (fig. 1 A) ; each oogonium 
arising from a separate peripheral cell of a node. Mature oogonia (fig. 1 A, E) 
ovoid-cylindrical, 760-940/* long. 6O0-7tSG •* wide; corona about 75 /t high, 225 /j. 
broad, of 5 small mucronatc cells, thickened at the apices ; spiral cells showing 
6 or 7 convolutions, each cell encircling the oospore -slightly more thaii once. 
Oospore black, cylindrical-oblong when ripe, 490-560/* long, 310-390/* broad, 
showing 4 or 5 ridges (fig. 1 E, F). 

* Botany Department, University or Adelaide. 
Traii>. Koy- $h£ S. Aust., 71, (2), 1 nwember, 1947 


Fig. 1 Profochara australis n. ?p, 
A, Apex of branch of a female plant, showing branch arrangement and position 
of oogonia. E, Antheridial plant showing general habit (natural size). C, 
Appearance oi nodal cells with a branchlet removed, the main stem being 
vertical, and with longitudinal rows of chloroplasts in the cells- D, Terminal 
mucronate cell and snbterminal inflated cell of a branchlet, showing the peripheral 
cells of the nodes. F. Mature oogonium attached to peripheral cell of a node. 
F, Oospore with spiral ridges fnot quite mature). G, Section of node of a 
branchlet, showing arrangement of central and peripheral cells. H, Antheridium. 
p.c, peripheral cells of node. (All drawings by camera lucida.) 



Fig. 2 Protochara iufiata (Filarski and Allen) comb. nov. 

A, Part of a plant showing the whorls of branchlets, together with oogonia and 
antheridin. B, End segment of a branchlet, showing the rnucro and basal cells 
of the node. C, Mature oogonium. D, Misshapen oogonium. E, Anthcridium. 

(After Filarski.) 

Antheridia borne similarly to the oogonia, ocioscutate, 800-1,150^- in 

As far as could be ascertained from limited material, the structure and 
development of vegetative parts and sexual organs agrees with that described for 
the Characeae by Fritsch (1935). 

Habitat — In swampy areas of shallow water (10-40 cm. deep) on top of the 
peneplain of the "breakaway" country between Mingcnew (about 15 miles from 
Mingenew) and the Irwin River coal seam, south-cast of Geraldton, Western 


Collected— 28 August 1947. 

From the same locality Lamproiimmnion macro pogon (Braun) Gphel comb. 
now/ 1 * and Nitella gclaitnosa Braun were collected. 

The type specimen (No. A 5,917 a) has been deposited in the herbarium of 
the Botany Department, University of Adelaide. Cotype specimens have been 
sent 10 the Herbarium of the Botany Department. University of Western Aus- 
tralia, aud to the Herbarium of the Royal Botanic Gardens, Kew, England. 

Protociiara infi>ata (Filarski and Allen) comb. nov. 

The following description of this specie* (as NUcltnpsis infiata) i-s give" 
by Filarski. 

Flantulae monoicac omniuo ccorticatae. Folia in verlieillis 4-6, uniarticulata, 
tegmentum ultimuin bicellulare, cellula ejus innma ut articulus unicus maxime 
infiata, ellipsoidea aut globosa-doliiformis, articulo aequilonga, cacumine mucrone 
minima acuta praedita. Foliola in nodo folii unico paucicellulati inevoluta, aut 
iu tiodo numeris 3-4 evoluta, papillaeformia, minima ventralia (antcriora). 
Corona stipularis in fundo vci'ticillorutn ioliorum nusqtiatn evolula. Oogonia 
solitaria aut bina nodo folii et basi foliorum orta; coronula oogonii e cellularum 
majorum verticillo pentamcro unico; antheridia oogorns majora solitaria ayt 
pluralia, hypngyna, in verticillis superioribus plerumque tantum modo evoluta. 
Flantulae propter folia maxime tumida, infiata, habitu exirnie differunt a 
Characcis ceteris; tantum Nitellam praeclaram Groves et Steph., plantulas Afri- 
canas juxta oppidnm <Cape Town> crescenres quodammodo in mentem 
revocant praecipue propter cellukts nonnulas internodii foliorum mterdum 
similiter valde tumidas, fere globosas, scd folia loco mucronis corona minutissima 
loliolis 3 composita praedito). 

Habitat — The following locality notes, given by Miss N. T. Burbidge, apply 
to both P, infla-ta and Char'ma ve.rlic'dktta (see later). 

The specimens were collected from shallow water in Lake Parkeyerring, 
about 5 miles south of VVagin, Western Australia. The water of the lake is 
brackish, especially hi a dry season. Collection dates- given by Filarski arc X. 1933 
for P. infiata and 5-6-35? for Charina verticiUata, Oa later visits Miss Burbidgc 
was unable to find further specimens. 

Filarski's figures of this species are reproduced as outline tracings in fig. 2. 
Details of nodal cells and antheridia are very indistinct in the original figure. 

The habit of P. infiata is very like that of P. australis, as will be seen from 
a comparison of fig, 1 and 2. Both species show simple, whorled branchlets, large, 
often inflated cells, and complete absence of stipulodes, bract-cells and bracteoles. 
The position of the sex organs is similar in both species, and neither species shows 
any development of cortical cells. The species may be distinguished as follows ; 

1. P. auslralis is dioecious, P. injlata monoecious. 

2. Coronal cells of the oogonium are small in P, australtSj large and con- 
spicuous in P. infiata. P. australis shows 6 or 7 convolutions, of the spiral 
cells, P. infiata 14 or 15. 

3. The sub-terminal iutcruodal cells of the branchlets of P_ australis show 
more pronounced asymmetry, and the lower cells are less Infiared than in 
P. infiata. 

Filarski states in his description of P. infiata that the antheridia are situated 
below the oogonia, but his figures (sec fig. 2 A) show some below and seme along- 
side the oogonia. He also describes the ultimate branch segments as hicellular. the 

( J > See Ovhcl: Notes on the Genera Lychnothamnus and Lamprcthsmniuut. Trans-. 
Km*. Soe, S. Aust. r 71. (2). 316. 


terminal cell being small and mucronate. In P. australis a distinct ring of nodal 
cells occurs at the base of the terminal mucronate cell, and it seems possible, in 
view of the close relationship between the two species, that close examination 
may show this to be the case in P. inflata. 

Why Filarski placed his plant in the genu* Nilellopsis is not clear. The one 
species of Nilellopsis (N. ohUisa J. Groves, from Europe) is distinguished by the 
presence of very long-, thick "bract-cells" which arise from the nodes of the 
branchlets and are almost as large as the ultimate hranchlet segment (Groves and 
Bullock-Webster 1924). "Starch stars," large, thickened, star-shaped nodes on 
the rhizoids are also characteristic of A 7 *, ohtusa. The absence of bract-cells places 
both the Western Australian species in a distinct genus from Nhellopsis. 

The corona of 5 cells surmounting the oogonium places Protochara in the 
tribe Chareae, as distinct from the Nitelleae which have a corona of 10 cells. It 
is necessary, however,, to modify the description of the Chareae given by Groves 
and Bullock-Webster (1924) and Groves and Allen (1934), in that the branchlets 
usually produce bract-cells at their nodes, but not in Protochara. 

The relationship of the genera of the Chareae, including Protochara, is given 
in the following synopsis (modified from Groves and Bullock- Webster). 

1. Stipulates and bractcoks absent. 

2. Bract cells absent Branchiets of 3 or 4 segments • Protochara n. gen. 

2. Bract-cells 1-2, very long- Branchlets of 2-3 very long 

segments .-, - IVitetlopsh Hy. 

1. Stipulodcs always present, sometimes rudimentary, Branch- 
lets simple, of 4 or more segxnents. Bract-cells normally 4 
or mure. 

3. Oogonia and atitberidia produced from separate peripheral 
cells ot the node C 1 ) {uc t , situated side by side). Stem 

corticate - -• "*• Lychnothavrnvs 

3, Oogonia and anthcridia produced from the same peripheral 
cell of the node. 
4, Oogonium normally situated Ifaelow the anthcridiurn 

Stem ccorticate Lamt>rothamnium 

J. Groves 
4. Oogonia situated above the aniheridium. Stem corti- 
cate or ecorticatc .,, , Chora L. 

The genera of the Chareae form an evolutionary sequence with Protochara 
as the most primitive, showing" an advance in vegetative construction through 
Nitellopsis, where only bract cells occur, Lo Lamp rot hamnhtnt-, Lychnoihamnus 
(which possesses a rudimentary cortex) and Chora, where bract-cells, bracfeoles 
and stipulodcs occur. The Diplostcphanae-tripiostichae section of Chafa repre- 
sents the culmination of the evolutionary series, showing complete 3-ranked 
cortication and two well developed whorls o( stipulodcs. 

The tribe Nitelleae is best considered as a separate evolutionary series 
parallel to the Chareae. No genus so far described provides a satisfactory link 
between the two tribes. 

Besides the absence of stipulodcs, bract-cells and bracteoles, Protochara 
shows another primitive character in the small number of convolutions (5 or 6) 
of the spiral cells of the oogonium of P. australis. The oogonium or P. inflata, 
however, shows 14 or 15 revolutions, and must be considered less primitive than 
P. australis. According to Groves and Bullock-Webster (1924) Nitellopsis 

< 3 > Sec Otthfcl, (oil ciU 

obtusa shows about 9 convolutions, whereas most species of Chara show more 
than 12. Fossil oogonia attributed to Characeae, judging from figures given by 
Groves and Bullock- Webster, commonly show a relatively small number of con* 
volutions (as low as 5 or 6). 

The relatively few segments to the branchlets, and the general simplicity of 
the thallus, also point to the primitive nature of Protochara. 

The naming of most fossil Characeous remains (usually oogonia), as species 
of Chara, makes Chara in this sense a very much wider genus than the Chara 
of living species. It is quite likely that fossil remnants would prove tu belong to 
Protochara if thallus structure were better preserved. 

It is evident that Protochara is the most primitive genus yet described of 
present-day Characeae, and appears to be more primitive than any genus of the 
Nitelleae. The general habit of the plant, however, consists of the stem with 
whorled branchlets that is so typical of the group as a whole, and the oogotiia 
and anlheridia are of the highly specialized type common to all species. The 
relationship and position of the Characeae amongst other plants remains as 
obscure as ever. 


Plantulae omuino ecorticatae. Stipulodae, bractae-cellulae et bracteolae 
omnino absentia. Monoecae ant dioecae, oogouia et antheridia a nodi pcri- 
mctrotis cellulis orta. 5 cellularum corona. 

Protochara australis n, sp. 
Plautulae dioecae, omnino ecorticatae, usque ad 10 cm. aha, subvirjdes, non 
incmstatae ; radiculae ramuhs et multicellulariae. Cautis crassus, simples aut 
ramulorum verticillis 4-7; internodia caulis 1-3 cm. longa, 0*9-l'5 mm. lata. 
Ramulae crassae pauco mcurvatae, in verticillis 4-7, quisque articulorum 3-4; 
segmentum mucroniformis ultimum longnm 110-200 /a, perimetrotis nodi cellulis 
S-6 in f undo ; segmentum subultimum magnum non aequab'ter inflatum ; segmenta 
inferiora longa 4-1 cm., lata -J-l-J- mm. Nodi perimctrotis cellulis 10-12. Stipu- 
lodae, bractae-cellulae et bracteolae omnino absentia. 

Oogonia in superiorum verticilloftun ramulorum axillibus verticillata, soli- 
taria aut bina nodis ramulorum; a perimctrotis cellulis propriis orta. Oogonia 
oviformia-cylindriformia longa 760-940 ^t T lata 600-780/*.; corona oogonii alta 
circiter 75 /*, lata 225^, composita cellularum S parvarum mucronif ormium ; 
cellulae spirilles convolutis 6-7. Oospora nigra, cylindriformia-oblongata, longa 
490-560 ^ lata 310-390/*, rugis 4-5. 

Aniheridia modo oogoniorum simili orta, octoscurata, diam, 1,150 /a. 


In the same paper as Nitellopsis inflata was described, Filarski founded ;i 
genus Charina. based on the one species C\ verticillata Fil. and Allen, from 
Wagin, Western Australia. None of (he stweimens on which the genus was 
named was fertile. 

FiJarski's reasons for founding a new genus are not clear, but were appar- 
ently based on the* vegetative form of the plant. His figures show a slender, 
verticillately branched plant bearing numerous whorled, 2-cclled dactyls. The 
occurrence of two- (sometimes more, rarely one) celled dactyls such as these en 
the branchlets is a characteristic feature of the Nitelleae, as distinct from the 


Chareae which hear only one-celled bracteoles on the branchlets. Filarski laid 
considerable stress on the apparent dimorphism shown by different branches (or 
plants?) of Charina; some branches bore only one-celled, blunt-ended, dactyls 
instead of the two-celled, mucronatc type. A feature of some, species of Nitella, 
however, is that the terminal cell of the dactyl is often deciduous, and this is 
probably the reason for the apparent dimorphism stressed by Filarski, 

From the figures given, the general appearance of the plant and its vegeta- 
tive construction offer no feature to exclude it from the genus Nitella, In fact, 
Filarski adds a note that J. Groves had suggested the plant was close to Nitella 
sublUUssima Eraun, and in the absence of fruiting material the naming of a new 
genus was hardly justified. Fertile material must be collected to prove whether 
Charina can be retained or not, and it is to be hoped that both this species and 
Protochara inflata will be rediscovered by Western Australian collectors. 

The authors are indebted to Miss N. T. Burbidge for information on the 
locality where she collected P. inilata and Charina verticillata. Mr. G. G. Smith, 
Department of Botany, Western Australia, also collected some of the material 
of P. australis. 

Fiiarski, Naotor 1937 "Idegenfoldi Charafelek Hatarozasa. (Determmatio 
Characearum Exoticarum)" Matematikai es Termeszettudomanyi 
Ertcsito, 55, 476-495. (Budapest) 
Fritsch, F. % 1935 "The Structure and Reproduction of the Algae," 1 
Groves H- J., and Allen, G. O. 1934 "A Review of the Queensland Charo- 

phyta." Proc. Roy. Soc. Qld., 46, 34-59 
Groves J., and Bullock-Webster, G R. "The British Charophyta." h (1920), 
' 2, (1924) 


By I. L. Ophel 


The genus Lychnothamnus was established by Leonhardi in Lotos XIII, 1863, p. 72, having 
previously been differentiated in 1845 by Ruprecht as a sub-genus to include the species in Braun's 
section "Charae pleurogynae," viz. Charabarbata Meyen, C. papulosa Wallroth and C. macropogon 
Braun. In "Fragmente einer Monographic der Characeen" 1882, Braun and Nordstedt recognised 
the genus as distinct, but C. papulosa (under the synonym of C. alopecuroides Braun) was removed 
into a new genus Lamprothamnus (later changed to Lamprothamnium by Groves, (1916), and C. 
stelligera Reich was added to Lychnothamnus, so that the latter genus consisted of three species: L. 
stelliger Braun, L. macropogon Braun and L. barbatus Leonhardi. 




By I. L. Ophel 

| Read 13 November 1947J 

,-, The _g entl s Lychnothamnus was established by Leonhardi in Lotos X1JI, 
lb63 f [>. 72. having previously been differentiated in 1845 by Ruprecht as a sub- 
genus to include the species in Braun's section i( Charae pleurogvnac/ 1 pig Chara 
harbata Meyen, C papulosa Wallroth and C. macropogon Braun In "Frag- 
mente emer Monographic c!er Characeen M 1882, Bwum and Nordstedt recognised 
the genus as distinct, but C. papulosa (under the synonym of C. alopcatrohh-s 
Braun) was removed into a new genus Lamprothamnus [later changed to 
Lamprothammum by Groves, (1916)]. and C. sielUqwa Keich was added to 
Lychnothamnus, so that the latter germs consisted of three species: L. stctliwr 
Braun. L. macropogon Braun and L. barbatus Leonhardi. 

As Groves (1919) points out, the distinguishing character of the gemi=. 
Lychnothmmtus as established by Leonhardi is lhat the antheridia arc produci-d 
by the side of the oogonium. Of the three species placed in the genus by Braun 
and JSordstedt it is only m the original tjpe (Lvchnothainnus barbatus) that the 
relative position of the sex organs can be satisfactorily ascertained, since 
Lychnothamnus stelhgcr (now A'iteltopsis obtusa J. Groves) te dioecious- and 
in Lychnothamnus macropogon, while the antheridm are normally produced at the 
branchlet nodes, the oogoma are almost invariably situated at the base and when 
situated at a branchlet node, scarcely ever occur at a node where there as also 
an anthendium present. 

Hy. in 1889, removed Lychnothamnus stitUgfr to a new trenus NUellopsis 
naming the plant NiMJopsis stclU v em f his new genus being bashed on vegetative 
characters. Considering the stale of knowledge at that time the action was 
hardly justifiable, since Lychnothamnus stdliyev satisfied Leonhardi's eeneric 
description to as great an extent as did Lychnothamnus macropogon However 
since it is now known that the relative position of the sex oralis is somewhat 
variable, it does seem that something more is needed on which to base generic 
distinctions. " * 

The removal of Lychnothamnus stcllu/cr left two species in the efcnite 
Lychnothamnus barhatus Leonhardi and Lychnolhamnus macropogon Braun. 

Groves (1919) examined specimens of Lychnothamnus mttcropaqon from 
Australia and Hongkong, lie found in specimens from Victoria one instance in 
which an oogonium and anthcridinm occurred together at a free node and in 
this case the two were produced side by side but proceeding from the same 
peripheral cell, corresponding to Lamprothamnium Groves ( Lamprothatmms 
Braun); in Lychnolhamnus barbatus, however, the antheridia and oogonia arc 
produced from different peripheral cells of the branchlet node, la the specimens 
from Hongkong two instances were observed in which an oogonium and 
anthendium occurred together; in both cases the two proceeded from the same 
peripheral cell, but the antherldium occurred below the oogonium in each case 
as in the genus Chara Linn. 

Despite the extremely close resemblance of the Vegetative parts of Lychno- 
thamnus macropogon to those of Lamproihamnium papulosum L Groves (so 
much so that some sterile fornis of both species would be iiKiisunguishable— 

Tra«i, InVj v» v . 5. Auat., 71, (2), 1 rircenitter, 1047 


Fig. 1 
A, B, C, Lamprothamnium mncropogon (Braun) comb, nov. 

A, Portion o£ sterile plant from Harriet River, Kangaroo Island (nat. size). 

B, Base of whorl, showing stipulodcs ; plant from Robe, X c, 30. C, Branchlct 
node, showing side-by-sidc arrangement of antheridium and oogonium. Plant 

from Harriet Rivtr. X c, 60. 

D, E, Lanipratham-nium papidcsum J. Groves 

(D, after Migula; E, after Groves and Bullock-Wcbster.) 

D, Portion of plant, nat. size. K, Base of whorl, showing stipulodes, Xc. 20. 

A, B, Lamprothamnium papillosum J. Groves (after Groves) 
A, Oogonium. X c. 60 B, Upper part of the branchlet with fertile node. Xc.20. 

C, D, E, Lychnothmnnus ba^rbatus, Leonhardi 

C, Base of whorl, showing stipulodes and rudimentary cortex. X 10. D, Fertile 

branchlet. X5. E, Oogonium, X 45. 


fig. 1 and 2), and that the relative position of the oogonium and anthemlvtwi 
appeared variable, Groves rejected the inclusion of the species in the genus 
Lamprothamniitm on the. grounds lhat it would mean abandoning the one dis- 
tinguishing character of this genus. He tentatively proposed the return of the 
.species to Chara. In "Review of Queensland Charophyta" (1934), II. Groves 
and Allen list the species as Chara macro pogon Braun. 

In all the South Australian specimens so far examined by the author only 
in two have an oogonium and anthcridium been found together. In one, from 
Kinchina (Coll, No. 25A), the anthcridium was situated below the oogonium. 
The other case was in a specimen from Harriet River. Kangaroo Island (Coll. 
No. 31), and in this case the two were side by side. In both instances the two 
reproductive organs were derived from the same peripheral cell. Another 
specimen from Kinchina (Coll. No. 25 B) had two antheridia situated at a free 
branchlct node, one above the other, and both proceeding from the same peripheral 

This derivation of the sex organs Irom the same peripheral celt confirms 
Groves' observations and supports his removal of Lychnothamnus macropagon 
from the genus Lychnothommts. Tt could not be said, however, that the above 
facts support his proposal of reluming ii in the germs Chara. 

Groves' reasons for ignoring the obvious relationship between Lamprotham- 
nimn papillosum and Lychnofhammts macrupogon are not clear. Uraunand Nord- 
stedt (1882) had remarked on their similarity, and Groves himself (1919) says, 
"In examining L. macropagon one is struck by the great similarity of its vegetative 
parts to our European Lamprothamnhtm papillosum/' 

This similarity extends to all characters of the plants (see also fig. 1 and 2). 
Both species are entirely ccorticatc with elongated stipulodes, usually one opposite 
each branchlct, at the stem nodes. The upper (usually fertile) whorls are con- 
tracted into heads, the branchlets in both species being incurved and of a small 
number of segments (3-4-5) bearing usually five bract cells at the lower -todes 
with hracteoles usually absent. 

Specimens of Lamprothamuium papillosum from South Africa (McNieol 
(1907) betray an even closer relationship by the production of a second row of 
stipulodes above the whorl of branchlets (common in South Australian specimens 
of Lye knot hamnus macro pagan), in the swollen segments of the luanchlets of 
the sterile whorls, and in the production of smooth spherical white root bulbils 
(seen in most South Australian collections;. The production of these bulbils by 
Lyehnothanmns macropagon is also mentioned by H. Groves and Allen (1934). 

Groves (1919) chose to ignore these similarities rather than abandon what he 
considered the one distinguishing character of Lamprathamniii}n< — that of the 
oogonium beiow the anthcridium — yet in his description of Lamprathommmn 
papillosum he states (Groves 1924, 2, 8) that the relative positions of 
the two are variable. In any case, instances of oogonia and antheridia 
occurring together in Lythnothamnus macropagon are so rare as to be regarded 
as atypical, and it would be hard to justify the use of these isolated instances as 
an ultimate basis of classification. Rather, the extremely close morphological 
resemblance of Lvchnothamuits 7Hacropogon to Laynprothamniuin paptdosum is 
a much more reliable indication as to the relationship of the Australian species. 

The Tiarne of this species now become::. Lam prof hamnium macropagon Braun 
comb. nov. with the following synonomy. 

Chara macro pogon Braun, 1843 

Lychnoiluimnus macropagon BraUft, 186S 

Macropagon ctustraikttm Migula 1891 


The amended generic description is as follows; — 

Lamfrothamxicm Groves, 1916 

(Lamprothamnus Braun, 1882, non Iliern) 

Stem and branchlets ecorticate. Branchlets of icw segments with usually 
five bract cells at a node. Stipulodes, in normal forms., long and declining, 
pointed and opposite the base of each branchlct; occasionally secondary ones 
produced. Sometimes a second series of slipulodes produced above whorl of 

Monoecious. Upper fertile verticils contracted into compact heads. 
Oogonium derived from same peripheral cell as antheridium when both occur at 
same branchlct node. Oogonium situated above, by the side of, or below 
antheridium, Root nodes producing spherical starch bulbils. 

Two species: — 

(1) Anthcridia and oogonia borne at same branchlet node — Lampro- 
thamnium papillosum Groves. 

(2) Oogonium at base of branchlets, anthcridia at branchlct nodes — 
Lamprothamnium macropogon (Braun) comb, nov. 

Three other species of Lamprothamnus (Lamproihavnmum) have been 
described. The most important of these is Lamprothamnus hanscnii Sonder, 
figured and described by Migula (1900). This species is related to Lamprotfumt- 
nium papillosum but differs in not producing the long dense fruiting heads, in 
having stouter branchlets and broader oogonia. This species is also closely 
related to Chora succina/a Braun, and seems to represent a well-defined inter- 
mediate form between Chara succiuata and Laniprolhamnimn papillosum. If the 
modified generic description of Lamprolhamnhim is applied, then this species, 
because of the stated differences, is ineligible and becomes Chara hanscnii 
(Sonder) comb. nov. 

The two other described species are considered by Groves (1924) to be 
extreme forms of Lamprothamnium papulosum. 

From the above discussion is can be seen that the typical species of Lampro- 
thamnium are connected by intermediate forms with species of Chara and the 
observations cast some doubt on the validity of the genus Lamprothamnium. 
Classification of the tribe Charcae, with the emphasis not on the relative position 
of anthcridia and oogonium but on their derivation would be more satisfactory. 
In this case the tribe falls into two sections : — 

(1) Antheridium and oogonium derived from separate peripheral celts 
of the branchlet node, (Nitellopsh and Lychnoihamnus).^ 

(2) Antheridium and oogonium derived from the same peripheral cell. 
(Lamprothamnium and Chara.) t 


The systematic position of the Australian Lychnothamnus macropogon Braun 
(Chara macropogon Braun) is reviewed. Figures are given of this and related 
species. From the evidence it is seen that the Australian species is more nearly 
related to Lamprothamnium papillosum Groves, hitherto the sole member of its 
genus, to which L. macropogon is now transferred. 

However, the validity of Lmnprothamnium as a genus is questioned. 

I 1 ) Also Protochara, see Womersley and Opbel* p. 311, this Journal. 



The author is indebted to the following botanists at the University of 
Adelaide:— Miss C. M. Eardley, B.Sc,, Systematic Botanist, and Mr. H. B. S. 
Womersley, M.Sc,, Lecturer in Botany, for advice and for the loan of specimens ; 
and to Mr. R. Perry, B.Sc, for specimens. 


Braun, A., and Nordstedt, O. 1882 "Fragmente einer Monographic der 

Characeen." Berlin. 
Groves, J. 1916 "On the Name Lamprothamnus, Braun." Jour. Bot, 54, 336 
Groves, J. 1919 "Notes on Lychnothamnitsf Jour, Bot., 57, 125 

Groves, J. 1924 (with G. R. Bullock-Webster). ''The British Charophyta." 
Ray. Soc, London 

Groves, H. 1934 (completed by G. O. Allen). "Review of the Queensland 
Charophyta." Proe. Roy. Soc. Queensland, 46, 34 

McNicol, Mary 1907 "The Bulbils and Pro-embryo of Lamprothamnus 
olopecuroides A. Braun." Ann. of Bot,, 21, 61 

Migula, W. 1900 "Die Characeen/' in Rabenhorst, Kryptogamenflora. 



By T. Harvey Johnston and Anne C. Beckwith 


The present paper deals with two new furcocercariae which have been rarely met with by us. One, 
Cercaria ancyli, belongs to the Strigeids, while the other, C. lophosoma, is a lophocercaria whose 
unknown adult probably is a blood fluke inhabiting one of the species of fish occurring in the 
Murray River. 




By T. Kar\i-.v Johnston and Anne C Beckwitk* 

(Fig. 1-8) 

[Read 13 November 1947] 

The present paper deals with two new furcocereariae which have been rarely 
met with by us. One, Cercaria ancyti, belongs to the Strigeids, while the other, 
C. lophosoma, is a lophocercaria whose unknown adult probably is a blood duke 
Inhabiting one of the species of fish occurring in the Murray River. 

We desire to acknowledge our indebtedness to Mr. G. G. jaensch and his 
family for help during our visits to Taileni Bend; and to the Commonwealth 
Research Grant for financial assistance. Part XI of the series is being published 
in the Records of the South Australian Museum, 1947 r 563-584. 

Cercaria ancyli n. sp. 
(Fig. 1-3) 

A very small furcocercarin, Cercaria ancyti, has been found parasitising two 
different hosts, the gastropod Amerianna pyramidata, and the freshwater limpet, 
Ancyhu australiius. It is the first occasion on which we have found l&fyal 
trematodes in the latter mollusc. The cercaria was first observed as an infection 
of one out of six Ancylus collected in the River Murray swamps at Tailem Bend 
in April 1947. In May 1947 three out of 154 Amerianna from these swamps 
emitted the same kind of cercaria. Jt is possible that, owing to the superficial 
resemblance of this species to C. angclae Johnston and Simpson 1944, the former 
mav have been collected on earlier occasions, but confused with the latter parasite. 

The cercariae are very active and swim, tail first, almost constantly. They 
live for about 36 hours at'room temperature, but keeping then! in a refrigerator, 
at about 1° Centigrade, in a vessel of water, prolongs the life to two or three day*, 
a use fid expedient, as material was scarce. 

Ten cercariae from each host, preserved in the usual manner, were measured 
with an ocular micrometer from a water mount. The measurements of the two 
lots of cercariae differ slightly, particularly in the proportions of tail-stem length 
co body length; but both body and tail-stem are highly contractile, and a* the 
cercariae possess no noticeable anatomical differences, the differences in the 
measurements may be accounted for by their development in different hosts. 

Measurements are given in micra and indicate the average, and (in brackets) 
the range. Cercariae Trom Ancylus*.— body length, 114 (81-144) ; body breadth. 
33 (27-41) ; tail-stem length, 112 (90-127) ; tail-stem breadth, 36 (27-41) ; furca 
length, 114 (99-124); furca breadth, 21 (18-25); anterior organ length, 34 
(25-43); anterior organ breadth, 22 (18-25); ventral sucker length, 18*7 (18- 
21); ventral sucker breadth, 17 (14-19). Cercariae from Amerianna:— body 
length, 127 (93-153) : body breadth, 37 (30-35) ; tail-stem length, 8S (72-100) ; 
tail-Mem breadth, 35 (28-45); furca length, 116 (95-139); furca breadth, 21 
(19-23); anterior organ length, 32 (27-37); anterior organ breadth, 25-5 (25- 
27) ; ventral sucker length, 187 (18-21) ; ventral sucker breadth, 19 (18-21). 

The spines are restricted to the anterior organ and ventral sucker. In front 
of the mouth are three rows of forwardly directed spmes, ten to twelve in all 

♦Zoology Department, University of Adelaide. 
Trnns. !<*>'. .<««:. S- Au*t. s 71, (2>, 1 December, 1947 


with usually only one or two in the most anterior row (fig. 1) and four to fi«! 
in the most posterior. A short spineless area succeeds the pre-oral spines, fol- 
lowed by a band of five or six irregular rows of spines around the front part «if 
the anterior organ. The portion of the anterior organ anterior to the first row 
of spines may be completely withdrawn, or pushed forward. The ventral sucker 
bears three rows of rather irregularly arranged spines, approximately 50 in 
number, and these, too, can be withdrawn into the cavity of the sucker. 

The digestive system consists of mouth, surrounded by the pear-shaped, 
highly contractile, anterior organ; a short pre-pharynx; well-developed pharynx; 
very short oesophagus ; and a short knob-like, caecum, all in the anterior third of 
rhe body. The caecum is at times partly lohulatcd in a manner suggestive of 
incipient division into two caeca (fig. 1). 

There is a group of six rather small penetration gland cells posterior to the 
ventral sucker, arranged in two group* overlapping anteriorly. The ducts of 
these pass forward and open each side of the mouth. There is a group of 
probably four pairs of very small "head-glands" in the region of the auteiior 
organ, which stain very deeply with neutral red used intra-vitam, and arc hence 
conspicuous features in such a preparation. The genital priinordium is a triangu- 
lar mass of undifferentiated cells between the bladder and the two sets of penetra- 
tion gland-ceils. The nervous system was not observe-d. A large, number of very 
small, highly refringent granules are scattered throughout the body. Whether 
these are part of the developing excretory system is uncertain. 

The stem of the longifurcate tail contains six pairs of caudal hodic*, the 
first rather smaller than the others. The usual stalked cells line the borders of 
the tail-stem. Both tail-stem and furcae are spineless Transverse and longi- 
tudinal muscle fibres are present. A unique feature of this cercaria is the curious 
cuticular thickening halfway along the furca, opposite the opening of the 
excretory canal. This is knob-like in some, in others mure spur-like, and is 
invariably present (fig. 2). 

The bladder is trilobed, consisting of a central portion with an anterolateral 
lobe on each side (fig, 1). Into this lateral lobe on either side opens the main 
collecting duct, which receives two secondary ducts at the level of the ventral 
sucker. The most anterior of these ducts drains the capillaries of two flame- 
cells, while the posterior, which is greatly coiled proximally, receives the 
capillaries of two further flame-cells in the body, and one in the tail at the 
level of the second pair of caudal bodies. Hence the excretory formula is 
2 t (2) -} (2 + (1) ) J = 10. In the region where the main ducts receive ihe 
secondaries a transverse commissure connects the two sides of the excretory 
system, passing across the body posterior to the ventral sucker From the 
posterior part of the bladder a wide duct leads hack, dividing to surround a small 
island of Cort, then continuing centrally along the tail-stem (tig. 3) and branch- 
ing at the base of the furcae into two vessels which open halfway along the 
furcae, opposite the cuticular "spurs." 

In one specimen a variation in the number of flame-cells on one side was 
observed. A third flame-cell in the hinder part of the body was connected wi(h 
the posterior secondary duct on one side only, making the formula for that side 
| (2) 4- (1+2+ (1) ) ] =6 (fig. I). This was apparently a precociously 
developed flame-cell of the mctacercarinl stage. 

The sporocysts (fig. 3) occur in the digestive gland in both hosts. They are 
slender tubular structures, usually tangled together in masses. Much of (he liver 


Fig. 1-3, C. ancyli 
\, body, showing spines, digestive system, glands, excretory system, ^ genital 
primordium; 2, tail; 3, sporocyst, showing cercaria emerging from tear in wall- 
Fig. 1 and 2 drawn from living specimens, outlines with camera lucida; 
fig. 3 from Canada balsam mount, also with camera lucida. 

tissue of the host may be destroyed. The living sporocyst is capable of slow 
waving movements. vSeveral sporocysts contained numbers of mature cercariae 
at the time when the first host (Ancylus) died, and some of the cercariae were 
observed pushing their way out through the walls of the sporocysts, but there is 


apparently no birth-pore. Characteristically, a number of constrictions divides 
each sporocyst into several lobes, and the end may be marked by a small knob. 
Length is very variable — a fairly long one measured 3 mm. There are no very 
marked differences between sporocysts from the two hosts, bet those from 
Amerianna tend to be slightly stouter and mure coiiexi than those from the smaller 
host, Ancyhts. 

Experimental Infections 
The second intermediate host of C. ancyli has not been ascertained. Attempts 
v/ere made to infect experimentally with the cercaria the gastropods Lymnaca 
lessoni, Amerianna pyramidata and Pfanorbis isingi; the fish Gamhusia affinis* 
and mosquito larvae; but results were in every case negative. 


As far as has been possible to ascertain, C. ancyli is the first cercaria ftom 
a fresh-water limpet to be described fully. Fielder, in WQ6, mentioned the^ presence 
of ccrcariae and of pigmented distome cysts in Ancylus tasmaniemis; while 
Cherry, m 1895, reported having seen ccrcariae from the same species, and in 
1917 from A. nustralictis. As these cercariae were not described, it US impossible 
to know whether any of them could have been C. ancyli The occurrence of this 
cercaria in Amerianna as well, indicates that its nearest relatives need not neces- 
sarily be parasites- of fresh-water limpets. 

One local cercaria. C. angdae, Johnston and Simpson 1944, also from 
Amerianna, resembles C. ancyli closely enough to make identification troublesome 
in routine examinations. The two species are distinguished, however, by a 
number of features, namely, size— C. angelae is considerably larger; number of 
gland -cells— eight in G\ angelae, six in C. ancyli; number of flame-cells— ten in 
C. ancyli, sixteen in C. angelae; and absence of "preacetabular bodies" in C. ancyh. 
They are also distinguished by the form of the alimentary canal, the spination, 
and ttf the fact that the tail excretory tubules in C. ancyli open half-way along 
the f tircac, but in C. angelae at the tips. 

Several well-defined groups of ccrcariae can be eliminated immediately from 
close relationship with C. ancyli, although possessing some features in common 
with the latter. The Elvae group of furcocercariae (Miller 1923) all possess a 
single pair of caudal flame-cells high in the tail-stein, apparently a group charac- 
teristic rather than a species characteristic, and common to several groups of 
furcocercariae; but they differ from C. ancyli in various fundamental features, 
such as the possession of a brevifurcatc tail. Probably much more closely related, 
though still distinct, is the Apaiemon group, again with one pair of flame-cells in 
the tail-stem, and also resembling our larva in size, general proportions and 
?.pination, but differing in having four pairs of gland-cells, seven pairs of flame- 
eelte, and well-developed caeca. C. mult icclfa fata Miller (1923) and its allies 
possess six gland cells posteriorly placed, but are quite distincL from C. ancyli as 
a group, as they have two pairs of flame-cells in the iail-stcm f a larger number 
of flame-cells in the body, and differ greatly in size, spine equipment and other 

The cercaria mostly closely resembling ours is C. dahema Cort and BrackeLt 
(1937), a parasite of Lymmea and Stagnicola, Of the relationships of this 
cercaria, Cort and Brackett said (p. 278) that they could find in the literature 
no other stri^eid larva resembling theirs. C\ ancyli has exactly the same excretory 
arrangement^ similar type of gut" and glandular equipment, similar pre-oral spines. 
six caudal bodies in the tailstem and somewhat similar body proportions, In 
actual size, however. C. dnhema is rather larger (body 156*4 long, tail-stem 179 # 


long, furca 196 /t long) ; it also differs in having spines back to the level of the 
pharynx, unpigjnented li eyespors," a pair of caudal bodies in the furcae. no head- 
glands and no f'urcal spur, and its sporocyst possesses a birth-pore. C. doheiwt 
is said to penetrate into a minnow (Funditlus sp.), and mctacercartae (un- 
described) recovered from the liver were considered to belong to that species, 
but investigations of the life-cycle were not completed. 

C ripom Bracket! (1939) resembles C. ancyli somewhat less closely. The 
glandular equipment is very similar, and the excretory system differs only in the 
presence of an additional pair of flame-cells in the posterior body; but it has no 
pre-oral spinas, more extensive bod> r spination, well-developed gut. eyespots and 
no bead-glands. C sine era Olivier (1941) has an excrctoty formula identical 
with that of C ancyli, and is very similar lu size,, hut has only two pairs of 
glands, a heavily spinose body, and long caeca. t\ gravida Miller (1927) and 
C. hirsuta Miller (192?) are two more species with exactly the same excretory 
arrangements as C\ ancyli; in C, granula there are also occasional variations in 
the number of flame-cells in the posterior part of the body, as in our larva. Both 
of Miller's cercariae have, in addition, very short caeca, and six caudal bodies, 
but differ from our larva in size, and markedly in their penetration gland equip- 
ment, although in both ibis is mainly posterior to the ventral sucker. Both have 
.setae on the tail-stem, 

Two olher cercariae with six pnst-acctabular gland-cells, C. higginsi Olivier 
(1942) and C. •<ixtilooui Olivier (1941), are quite distinct from cVancyli; both 
are considerably larger larvae titan ours, and both have sixteen flame-cells, two 
of which are in the tail-stem. 

Cercaria lophosoma n. sp. 
(Fig. 4-8) 

In the course of examination ot 2.920 specimens of ihe gastropod, Notopala 
hanlcyi, tor trcmatode infestation, * minute new Jophocercaria, Cercaria lopho- 
soma, has been recovered from two snails. These two infected specimens were 
collected in May 1045 and March 1946 respectively at Swan Reach on the River 
Murray; ten other collections of this mollusc made at the same place between 
April .1942 and March 1947, and two collections made at Keumark and Morgan 
respectively, yielded no further specimens infected with the same cercaria. It is 
possible that the cercariae at times escape notice because of their small size, 

C\ lophosoma is emitted mainly in the middle of the day. A few appear by 
10.30 a.m., but large numbers are not emitted until between 12 noon and 2.30 pm. 
When .sufficient numbers are present, they tend to form a swarm in the tube. 
They arc planktonic organisms and swim very little, hut will respond to vibration 
of their tube with a few jerky bending movements of the tail. They float in 
various positions (fig. 6). The length of life is about 48 hours. 

Cercariae. fixed by adding an equal quantity of boiling 10% formalin to the 
water in which they were swimming, were measured in a water mount with an 
ocular micrometer. The measurement of the breadth of the furcae excludes the 
fms, which in preserved material arc often bent or shrunk. The averages of ten 
measurements are given in micra, with the ranges in brackets: body length. 95 
(82-108) J body breadth ar widest part, 27 (21-32) ; tail-stem length, 196 (180- 
213) ; tail-stem breadth. 19 (16-21); furca length, 63 (54-73); furca breadth, 
7 1,5-SO ; anterior organ length, 18 (16-19) ; anterior organ breadth, 16 (14-19). 

A difficulty encountered in studying this cercaria was the impossibilitv of 
making a ventral mount during life, partly because of the lateral compression of 
the body, and partly because of the stiff cuticular crest. Hence the measurement 
Riven as "body breadth'* is more exactly "body depth" 

Fig. 4-8, C. laphnsomt 
4, body greatly contracted; 5, general features of body and tail, and excretory 
system; 6, various positions assumed when floating; 7, body in extended position, 
showing glands, central granular mass, and genital priniordium; 8, sporocyst. 
Fig, 4, 5, and 7 drawn from living specimens, outlines with camera lucida; 
fig. 6 drawn freehand from living specimens in a drop of water without covcrsiip; 
ii^- 8 drawn with camera lucida from Canada balsam mount. 



The body of the cercaria is, like that of C. helvetica x%ri (Dubois 1927, 
p, 27) highly contractile. There are five to six rows of fine straight spines round 
the front of the anterior organ. There are no further spines on the body, although 
when greatly contracted it has a spiny appearance, due to the intense wrinkling 
of the rather stiff cuticle (fig. 4). Small spines are scattered along the borders 
of the tail-stem, and more thickly on the fleshy part of the furcae, 

The anterior organ is highly contractile and, as in Sewell's Indian Lopho- 
cercariac (1922, p. 46), definitely snout-like. There are no "hollow, conical 
spines" on the tip of this snout, as described for several Lophocereariae, bat at 
times drops of secretion from the gland-ducts which open on the anterior surface 
of this organ, and which are highly refractive, may be seen. There is no ventral 
sucker. There ate no eyes, though they have been described for some Lopho- 

The glands (fig. 7) are numerous, mainly in the middle and posterior regions 
of the body. Neutral red and .Nile blue sulphate were used as intra -vitam stains, 
Delafteld's haematoxylin and acetic acid alum carmine for permanent prepara- 
tions. Two unicellular glands in tandem are situated ventrally, their ducts pass- 
ing forward to open on the ventral surface of the anterior organ, separate from 
the other ducts. These two glands stain deeply with both neutral red and Nile 
blue sulphate, while a large group of glands dorsal to, and extending posterior 
to them, stain more lightly with neutral red, but just as deeply with Nile blue 
sulphate. These glands could not be accurately counted, but number more than 
twelve. Their ducts pass forward together centrally, to open on the apex of the 
snout. In the middle part of the body is a group of deeply-staining granules 
surrounded by a granular mass. It could not be determined whether these were 
the rudiments of the ventral sucker, or part of the genital primordium. A 
triangular-shaped mass of cells, staining deeply with acid alum carmine and with 
Delarleld, situated just in front of the bladder, is certainly part of the genital 

No trace of a digestive system is present, not even a mouth-opening was 
observed. The nervous system could not be distinguished- 

The dorsal crest is a conspicuous feature of this cercaria. It is fine* trans- 
parent and slightly yellow, and is apparently formed of an extension of the cuticle. 
Its shape varies with the extension and contraction of the body (fig. 4 and 7). 

The long slender tail is, together with the furcae, two to three times as long 
as the body (fig. 5). It is provided with a few very fine, hair-like structures 
dorsally, as well as the minute spines mentioned above. There are numerous 
small rounded caudal bodies grouped round the central axis of the tail; these 
disintegrate readily under pressure. A lew scattered nuclei in the tail-stem stain 
deeply with neutral red in life. The tail-musculature is well developed. The 
principal fibres run obliquely in two direction?., and hence in two sets, one dorsal 
and one ventral ; when viewed from the side they appear to be arranged in hening- 
bone fashion, the apices of the one set directed forward, those of the other back- 
ward. Longitudinal fibres also are present, 

Kach short slender furca is provided with a fine cuticular flange or fin, 
extending round the whole furca. Longitudinal lniuele fibres are present. At 
the end of the furca the flange forms a pocket-like or flask-like fold, open 
posteriorly, as is characteristic for this type of cercaria. At the base of this 
pocket* on the tip of the furca, opens an excretory pore. 

As far as could be determined before the host died, the excretory system & 
of the usual pattern, i.e., 2 (2-r- ') = 6. The two sides of the excretory system, 


if it be bilateral, were, however, never seen simultaneously in one specimen. The 
bladder (fig. 5) is bilobed, and from each side a duct passes forward. From this 
point, presumably owing to the fact that a ventral view was never obtained, only 
one set of tubules could be seen. The main duct branches into two, one passing 
forward to receive the capillaries of two flame-cells, the other passing back to 
receive a single flame-cell at the level of the bladder. There are no flame-cells in 
the tail. There is a long island of Cort, from which a single duct passes back 
through the clusters of caudal bodies, to divide into two tubules about three- 
quarters of the way bade along the tail-stem (fig, 5). One tubule passes into 
each furca, opening at the tip into the flask-like extension of the flange. 

Experimental Infections 
Unsuccessful attempts have been made to infect the fish, Gambusia affinis 
and Carassms attraius, and a tortoise, Emydara tnasquarii, with the cercaria- 


The sporocysts are very small, round, or oval bodies (fig. 8), and at the time 
of examination, after the death of the host, contained only germ-balls, which 
stained more deeply with acid alum carmine than did the rest of the sporocyst. 
Large masses of these parthenitae were packed together in the liver. They vary 
somewhat in size, and also in shape, according to the pressure of the surrounding 
tissue. The average length of ten, measured with an ocular micrometer from a 
canada balsam mount of a stained fragment of liver, was 101 /a, varying between 
84,*and 120 /*. 


Tbe "Lophocerca" group of furcoecrcariac, so named by Liihe in 1909 to 
include C. cristata La Valette and C. microcristata Ereolani, was defined fully by 
Sewell in his "Cercariae Indicae" (1922); the essential features of cercariae of 
his "Lophocerca" group being the very small size, the relatively long, brevi furcate 
tail with furcac provided with a fm-fold; body with a crest; eyes (in many); 
gland cells in mid-body; anterior organ snout-like rather than sucker-like; no 
mouth, alimentary canal or ventral sucker; excretory formula, 2 (2 -J- l) = 6j 
development in small oval or rounded sporocysts. Scwcll himself added four 
cercariae to tins gtoup (Cercariae indkae fX t XIII, XXjClX, and LV), and since 
then a number of other Lophoccrcariae have been described. 

Odhncr (1911), Scheuring (1922), and Ejsmont (1925), have shown that 
certain forms belonging to this group develop in the blood of Cyprinid fish into 
species cf the genus Sanquinicolo, Wall (1939; 1940), on the other hand, 
described a cercaria possessing the characteristic dorsal crest, which developed 
into Spir orchis panmx Stutikard, a blood-fluke of American freshwater tortoises- 
rlowever, the life-history of another species of Spirorchls, S. elephaniis Cort, 
was described by Wall in 1941, and its cercaria possesses no dorsal crest. Thus.. 
if the possession of a dorsal crest be- the only characteristic necessary for inclusion 
of a cercaria in the Lophocemi group, it would at once become a highly artificial 
group, closely related forms being separated, and unrelated forms being grouped 
together. Under Sewell 's more limiting definition the presence of a dorsal ciest 
is only one of several distinguishing features, and the cercaria of Spirorclm 
parvus is excluded from the group by the possession of an alimentary canal 
f apharyngeal ) and a ventral sucker, and by its excretory formula of 
2 [ (1 + 1 -f 1) + ( 1 + 1 + (1) ) ] =12. Furthermore, Wall's description of 
me crest of the cercaria of 5*. p&r&frs indicates that its structure is somewhat 
different from that of the true Lopbocercariae. 


C. Ivphosoma is a Lophocercaria belonging io Se well's group, but distinct 
(mm all other members. In size it is closest to C helvetica XV t Dubois (1929), 
which, like our cercaria, has numerous gland cells and no eyes. Complete com- 
parison is not possible because Dubois 7 description is very brief; the host of 
Dubois' larva is a very different gastropod, however, «te 4 Lymnaca. C. lopho- 
soma differs from Swell's Cercaria indica IX (from " Indoplanorbis and 
Gymuhts), XXXfX (From Amnkola), and LV (from Awmcula), m having no 
hollow spines on the tip of the snout, no eyes (pigmented or oihcrwisc). a greater 
number of glands cells distributed differently, and finally, in size— heing slightly 
Inrger than C. indica IX, and somewhat smaller than cither C indica XXXIX or 
C. •indica LV. C. indica XIII (from Ammcola and MrUnioidex) has no 
apical spines, but is larger than C\ lophosonm, possesses non-pigiucuted eyespnts. 
fewer gland cells* no spines on the furcae, and a papilla-like structure ventrally, 
thought to he a rudimentary genital papilla. The body of Scheming';; C\ Sangidni- 
colae inermis (from Lymnaca) is slightly larger, according to Fjsmont's figures, 
and the tail is larger m proportion to the body than in our cercaria. The measure- 
ments Ejsmont gives for his C\ Sa^iithrkolae spp. from Bithynia and trout 
Lymnaea are both somewhat larger than those of C\ lophosoma} he shows two 
e*creu»ry canals in the tail-stem, and though the large number ot gland cells is 
suggestive of our cercaria, Ejsmont figures also a short, blindly-ending gut, in a 
somewhat similar position to the two special gland cells of C. lophosoma. 

C cristvtit La Valctte (1852). from Lymnaea, which Scwcll considered 
(p. 53) might be the same organism as his C. indica XIII, is a larger cercaria 
than ours, according to Ejsmont's figures; the cercaria which Wescnherg-Lund 
(1934) describe? as C. cn'stata has the same body measurement as C. hphosam^ 
but the tail-stem is longer and the furcae very much longer; there are special 
apical spines, two excretory canals in the tail-stem, and the sporocysts arc long 
and provided with a sucking disc. C. microcristata (from Bithynio) is, according 
to figure* given by Ejsmont, smaller than C. lopfwsotna and all other known 

Martin (1944) ledescribed a marine Lophoccrcaria, previously described by 
Linton (1915), and named C. kwssi by Stunkard (1929). This larva is remark- 
able tor having as its host an annelid worm, Ilydroides. In all other respects it 
is closely allied to the Lophocerca group, and the sporocysts are apparently of 
the characteristic type. Both Linton and Martin consider it closely related to 
cercariac of the genus Sanguinkola. It is, however, quite distinct from C. hpho- 
turna in being larger, but with turcac shorter, and in having twelve to thirteen 
rows of spines around the snout, and fewer glands, arranged differently. 

t-\ lophosoma is distinct from C. xewelli Faust (1926), from tturmtpia, which 
js larger, has -only two pairs of glands, a pair of partly-pigmentcd eyes, and as fa* 
as- can be seen from the figure, no fin-folds on the furcae. Three other dorsally- 
crested cercariae, C. whitenioni Croft 1933, C\ brevifurca McCoy (1926) from 
Planorbis, and C, bombaycmis No. 8 Soparkar (1921) from Planorlris and 
f.ymnaea, show somewhat dubious affinities with true Lophocercariac, because all 
possess simple, apharyngeal alimentary canals, and fin-less fnrcae, and they 
develop in rediae. C. bombayensis No. 8 and C. whitentoni possess also a rudi- 
mentary ventral sucker, and the excretory systems are more complex than that 
of the true Lophocerca type, It is possible that these three cercariae may be 
more closely related to Spirorcltis or perhaps to Clinostomunu since the cercaria 
of CHnoshnmm marginatum has a dorsal crest, finlcss furcae, alimentary canal, 
ventral sucker rudiment, and live pairs of flame cells, and develops in rediae 
KruU 1934) 



Cercaria ancyli n.?.p., a parasite of Ancylus australicus and Amerwnna 
pxramidata, is a longi furcate pharyngeal slrigeid distomc cercaria with six pene- 
tration glands, head glands, ten flame cells, an excretory commissure behind the 
ventral sucker, very short gut, and a furcal spur. Mclaccrcaria and life cycle 
arc unknown, 

Cercaria lophosoma n. sp., a parasite oC Notopala hanlcyi, is a dorsally- 
crested breviturcate, non-ocellatc cercaria with numerous unicellular glands, but 
without a ventral sucker or alimentary canal. It belongs to Sewell's group,, 
Lophocercatia. and is closely related to the cercaria of Sangtihdcola. 


rlRACKK'rr, 5. 1939 jour. Parasiiol,, 25, 263-266 
Cherry, T. 1895 Proc. Roy. Soc. Vict., 8, 183 

Ciilery, T. 1927 Bilbarziasis and the Danger of the Disease becoming En- 
demic in Australia. Commonwealth Defence Dept., Melbourne, 1-20 

C\na\ W. W., and Brackett, S. 1937 Jour. Parasitol., 23, 274-279 

Croft, J. W. 1933 Trans. Amer. Micr. Soc, 52, (3), 259 

DuiiOis, G. 1929 Bull. Soc. Neuch. Sci. Xat., 53, 27-28 

Ejsmoxt, L. 1925 Bull, de l'Acad. Polonaise Cracow CI. Se. Math. Nat., 877- 

Faust. E. C 1926 Parasitol., 18, 102-103 
Fielder, W- 1896 Vict. Naturalist, 12, 139-140 
Johnston, T. H., and Sku'son, fc R. 1944 Trans. Roy. Soc. S. AusL. 68, 

Ktn.TLL, VV. H. 1934 Proc. Helminth. Soc. Wash., 1, (2), 34-35 
Limtok, E. 1915 Biol, Bull., 28, 198-209 (nor available) 
LiJiiE., M. 1909 Parasitisehe PlaUwv.rmer, 1, Trcmatodes 
Martin, W* E. 1944 Trans. Amer- Micr. Soc. 63, &)\ 237-243 
McCoy, O. R. 1926 Jour. Parasitol., 15, 204-206 
Miller, II M. 1923 Jour. Parasitol., 10, 35-46 
Miller, IL M. 1927 Parasitol., 19, 72-74 
Odhner, T. 1911 Zool. Anz., 33, 33-45 (not available) 
Olivier, P. 1941 Trans. Amer. Micr. Soc, 60, 45-52 
Oltvter, L. 1942 Trans. Ameri. Micr. Soc, 61, (2), 168-179 
ScunuRiNG, E. 1923 Zool. Jahrb. Abt. f. Anat., 44, 265-310 (not available) 
Sewkll, R, P>. S. 1922 Tnd. Jour. Med. Res,, 10, SuppL 44-63 
Soparkar, M- B. 1921 Ind. Jour. Med. Re*., 9, 23-32 
Wall, L. D. 1939 Tour. Parasitol., 25, (6, SuppL), . 28 
Wall, L. D. 1940 Science, 92, (2,390), 362-363 
Wall, P. 1). 1941 Amer. Midi. Nat. 25, (2). 402-411 
Wesexberg-Lu.m>, C. 1934 Denk. Kgl. Dansk. Vidensk. Selsk. Skr. Natur. 

Math. Afd., 9 Rackke, 3, P8-103 



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i873 12 

ENDOWMENT FUND as at 30 September 1947 
(Capital— Stock, etc., Face Value, £6,042 I8s. 7d,; Cost, £6,041 8s. 7d.) 

£ s. d. £ s. d 
1946— October 1 
To Balance — 

Aust. Inscribed Stock 6,008 10 

Savings Bank of S.A. 17 3 7 

Bank of Australasia _ . 15 15 

6,041 8 7 

,, Interest — 

Inscribed Stock .... 20O 9 6 

Savings Bank of S.A, 1 13 8 

202 3 2 

i6.243 11 9 

1947— September 30 
By Revenue Account 
„ Balance — ■ 

Aust. Inscribed Stock 6,008 10 
Savings Bank of S.A. 32 18 7 

£ s. d. £ s. d. 
202 3 2 

6,041 8 7 

i6,243 11 9 

Audited and found correct. The Stock and Bank Balances have been verified by certificates 
from the respective institutions. 9 

O. GLASTONBURY, F.A.I.S., A.F.I.A. ) Hon. 
F. M. ANGEL J Auditors 

Adelaide, October 1947 


Hon. Treasurer 


1929 Prof. Walter IIowcbin, F.G.S. 

1930 John McC. Black, A.L.S, 

1931 Prof. Sir Doiiola.s Maw son, O.B.E., D.Sc, B.E., F.R.S 
1933 Prof. J. Burton Cleland, M.D. 

1935 Prof. T. Harvey Johnston, MA., D.Sc, 

1938 Prof. J. A. Prescott, D.Sc, F.A.T.C. 

1943 Herbkrt Womeksley, A.L.S., F.R.E.S. 

1944 Prof. J. G. Worm, D.Sc, Ph.D. 

1945 Cecil T., M.A., B.E., D.Sc, F.G.S. 

1946 Herbert M. TTale 



Those marked with an asterisk (*) have contributed papers published in the Society's 
Transactions. Those marked with a dagger (f) are Life Members. 

Any change in address or any other changes should be notified to the Secretary. 

Note— The publications ot the Society are not sent to those members whose subscriptions 

are in arrear. 

5, ate .- o( Honorary Fellows 


1945 *Black J, M., A.L.S., (Hon. causa), 82 Brouffham Place, North Adelafide— Few 

Medal, 1930; Fellow, 1907-45; Council, 1927-31; President, 1933-34; V ice-President, 

1945. *Fp.nner > C. A. E.. D.Sc. Alexandra Avenue. Rose Park. Adelaide— Fellow, 1917-45; 

Council, 1925-28; President 1930-31; Vice-President, 1928-30; Secretary, 1924-25; 

Treasurer, 1032-33; Editor, 1934-37. 


1945. Abhte, Prof. A. A., M.D., D.Sc, Ph.D., Umvers ; ty of Adelaide. 

1935. Adam. D. B., B.Agr.Sc. Wa : tc Institute (Private Mail "Rag), Adelaide— Council, 

1939-42; Vice-President, 1942; Librarian, 1942-. 
1027 *Al»erman A. R-. Ph.D.. D.Sc. F.G.S. Div. Tndus. Chemistry, C.S.I-R., Box 4331. 

G.P.O., Melbourne, Victoria— Caunnl 1937-42. 
1931. Andrew, Rev. J. R.. c/c 212 Voune Street. North Unley. 

1935 *Anj)REWArtha. H, G„ M,Affi\Sc D.Sc, Wake Institute (Private Mail Bag), Adelaide. 
1935. *A-nt>rkwavttta. Mrs. II. V., B.Affr.Sc. M.S., (nee liV. Steele), 29 Clarctnont 

Avenue, Netherhv, SA. 
1929. Ancel, F. M., 34 Fullarton "Road, Parks ide. S.A. 

1939. * Angel, Miss T,. M., M.Sc, c/o 2 Moore Street, Toorak, Adelaide. 
1945. Bartlett, H. K., L.Th., Btirra, S.A. 

1032. Bww, P. R., D.D.Sc. L.D.S.. Shell House, 170 North Terrace. Adelaide. 

1928. Best, R. J., M.Sc., F.A.C.I.. Wait** Institute (Private Mail Bag), Adelaide. 

1934, Black. E. C. M.B.. B.S., Maffill Road. Tranmere, Adelaide. 

1945. Bonython, C. W-j B.Sc-, A.A.CT., 2f>9 Domain Road. South Yarra, S.E. 1, Vict. 

1940. Bonython. Sir J, T^avtnc.ton, 263 Hast Terrace. Adelaide. 

1945. FrwwsMA, C D.« B.Sc.For., 2 Cehic Avenue. South Road Park, S.A. 

194S_ Brovguton, A. C. Mt. Scl'e Station, via Copley, S.A. 

1947. Howes, D. R.. B.Sc, 51 Eton Street. MaWn. 

1039. Brook man. Mr?. R. (nee A. Ilarvcv), B.A., Meadows, S.A. 

1944. BuKHinc.E. Miss N. T. r M,Sc, CS.LR. Div. Plant Industry, PO. Box 109, Canberra, 

1923. Rnimorv, R. S., D.Sc, University of Ar!ela : de. 
1922 ♦Campbell. T. D. t D.D.Sc. D.Sc. Dental Dept, Adelaide Hospital Adelaide— 

Council 1928-32. 1935, 1942 45; Vi<-c~Presidcni. 1932-34; President, 1934-35. 
1944 Casson P. B., B.Sc, For. (Add.), Dept. For., Mount Crawford Forest, S.A. 



1929. Christie, \V\, M.B., B.S., Education Department, Social Services nt Pirie Slices 

Addmik— Treasurer, 1W3-38* 
W& •CtgiJTO tittg J. B M.D., University of Adelaide-/ V m? tfc&i 1933; fowl, 
lfPli tr . JpWR Wg^g.; l*rthdqt* J 1927-28; 1940-41; FiwFresMenL 1920-27, 19-11-42. 
I93tt *CoLgmioi;.N, 1. I., M .Sc„ 10 French Street, NcUierbv, S.A.— Secretary 1942-43 

1907. *P^K ^'*I-;. U t -? t: -\ J AA - J Cr - University, Adeiaidc-tVw^, 1938-41, Ficc-Presidnu, 

1941-42, 19-13-44; Prestdrul. 1942-43. 
IW *CoowM t H. M., 51 Hastings Street Glaidsr, S.A. 
1044. Cokmsh, Melville, State Bank, Pirie Street, Adciaide. 
194?, * Con on, B. C, S.A. Mxieeuni, Adelaide— Ct-itnal 1<&N$ 

1924. » E Crkswgky. Sir C:. T. C, O.S.O., MIX, F.R.C.F., 210 North Tt-rrace Adelaide 
19.v. n.Rfn-KKit, R. L,, M.Sc., Waite Institute (Private Mail JW.h Adelaide- -S'ecrrtar™ 

1943-45; Ou^nY. 1945-47, * '" 

£&' T ,CK £ s Vr"' % ^HSfe Gtm - G&****. #** Department. Flinder.s Stmt, Adrlaicfc 

1930. DijKi EX V., Hospitals Department, Rmidle Street, Adelaide, S A 

194-1 Dvxstoke, S. M. JU M.B., B.S., 124 Payne-ham Road, St. Peters, Adelaide. 

193!. lKvver, j. St, M.B., l^S.. 11 Gstttfh Road. Kin^-wool, S.A. 

2933. W%u^y:%£& C. M;. B^h Wake Institute (Private Mat] BasO, Adelaide— CViHwrtf, 

1945. Euaionus, S. J., H.A.. M.Sc, 56 Ficbci Terrace, Mile End S A 
1002. ♦Ei.qvist, A. G M 19 Farrell Street, Gleneliy, S.A. 

US' J?* i0SS ' Miss ^ )*< &&». 8 Trior's Road, Mitcham, SA. 
1 ( ^2/. *FrxLAvsox,, IT. H.. 305 Ward Street, North Adelaide -Coinu-ii 1937-40 
Ittt ♦h'RV TT K. ; D.S.O., M.D„ U.S.. R.Sc. F.R.A.C.F., Tom. Hall, Ade'airie-Cm^aL. 
1933-.V; l'irc-} J rrsirf:i;f^ VJ37-38. 1939-40; Fresidmt l93*-39 

1932. *Giasox, K. S. IT. f KSc„ 297 Cross Roads. Clarence Gardens, Adelaide 

?*$?■ f<' LASr0XR ™ Y . J- O. G.. E.A., M.Sc., Dip.Fd, fietftir Terrace, Mumv ttfcfefe, S.A. 

£12' tOx-astonbl-ky, O. A.. Adelaide Cement Co., Grenfelf Street, Adelaide. " 

1927. Godfrey, F. K., Robert Street. Payncham, S.A. 

1935. fGoLiiMACK. H„ Coromandel Va^ey, S.A. 

1939. Goooe, J. R., R/Agc&q., P.O. ftox T80. Whvalla 5 A 

i&5? IS°P^ 5m J AxlE * tt.. Gilbert House, GiihWr Place, Adelaide. 

1910. *Grakt, Pkof. Sir Kf.kk, M.Sc., F.7.1\, Univers-rv of Adelaide. 

1930. Gray, J. T. f Orroroo, SA, 

1933. Greaves, H., Director, Botanic Gardens, Adelaide, 
3904. Griffith. R B.. Dnnrob'n Ihwd, Brighton. S.A, 

1944. Gif^'Y, D. J.. 13-Sc. Mineral Resources Survey. Canberra, A.C.T. 

1922. IHAlr. H. M. Director. S.A. Mu-ieum, Adcla'de- -Perra Mrdaf, W6; CtmneU ]-0Sl S4i 

Ficc-Pn-stfenr, 1934-3."\ 1937-3S; President, 1936 37; r^jfAivr; 1938- 
19*5. *TfA^r.v, 5J W - j. H., tl:cc A c. Beckwith), M.Sc., 59 GrccnhilJ Road, Toorak Gardrn^ 

1944. pasts, J. R.. i;,Sc.. 94 Archer SU'cct. North Adelaide, S.A. 

1W. Mi.:\nt'R50\ ( D. L f \V.. CntJ«;5^j»e, "Woodi'^rd, N.S.W. 

1944. Hfrriot, R. t, B.Agr.Sc. Soil Con.^rvator. Begfe of Atrricu'ture. S.A. 

Ki^l. *Hos«FKr.r.. P; S., M.Se,, 1X1 Fhhvr Sireft b'tstterOfh S*V 

1944. HtHrpfjtj D. S. W., 238 Pavneharn Road, P.T)nelnnv' S A. 

}^l T HT ' ms J- T - BSc - 15 Sl.-rbourne Road, Moc!indi e Gardens. S.A. 

1928. Ifouu., V., Knrraha, Burnside, S A. 

1942. Jexicjws, C. F. FL ; Department of Agriculture. St. Genre's Terrace, Perth W A 

WML *TeNNrsoN-, RRVi J. C., 7 Frew Street, F^larton, 

1945. *T'^sur, R. \V„ RSc. 3 Alma Road. Ftdlarton, S.A. 

I! } TJp9*$toj E- A. M.D.. M.R.C.S.. ,; Tarni W Port Noarhuma. S.A. 

1"'21. *John-sto.v, Prof. T. IL. M.A., D.Sc, of Adelaide— I'mo Mr<ioJ l^.V * 
di'H-tV, 1926-28, IMfrl f'ire-f'rcsidrvj, 1028-31: Present, 1931-32; Sr'rrctar" 
193S-40; Pep. f<iwa and Flw* Boned, 1932-39; Editor, 1943-45 

1939. tKhaichar, II. M-, Ph.D., M.E., F.R.G.S., Khakar Buildinffs, C.P. Tank Road, Rom- 
bay, India. 

1933. *Kr.?:FMAx. A. V/., M.Se., University of Adelaide; Srrrctan; 1945-, 

19-15, I.ayton\ E. K. T.. Co'onfal Setiar Remicrv Co,. Kinrr AViT-iam Street Adelaide. 

1922. L^MKtN, G. A., M.D., B.S.. F.R.C.P. AM.P. Bnildmft", Kinir Willia.n Street, Adelaide. 

WL LfPnRooK Mrs. W. V., (tiec N. H. Woods). M.A., Flimatta Street. Reid A-CT 

193*. Maddkrv. C. B.. U.D.S.. DTXSc, Shell House, Korth Terrace, Adelaide. 

1932. Mann, F. A. ( C/o Pank of Adelaide, Adelaide. 

1939. Marshall, T. J., M.A^r.Sc, Ph.D., Waite Institute (Pfivate Mmt Ban), Adelaide. 


Date of 

£ lection* 

1020. Marttv. V C„ MA, Technical Hi«h School, Theharton, SA 

21)05. *Mawson, Proi . SXH Donc.T.As, O.R.E., D.Sc., B.E, F.R.S.. University of Adelaide— 

Vm» Medal 1931, PscM-nt, 1024-25, 19444S; Vke-Prcudeitf, 1923-24. 1925-26; 

Council 1941-4.1 
l*J2tt. Mayo, The Hon. Mr. Jt'STrcK, L1..B,, KX., Supreme Court, Adelaide. 
194.3. McCarthy, Mtss D. f*., B.A., E,Sc. 70 TTalton Terrace, Kensington Park. 
1044. McGilp, L. K., Ramsgate. Hotel, Henley Beach, S.A. 

1944. *Mnxs. K. R., D.Sc. F.G.S M Mimes nci>artment f Flinders Street, Adelaide.. 
1439. MrNcrtAM, V H.. Hammond, SA. 

1^25. tMitchkll, Prof- Sir W., K.C.M.G., MA, D.Se., Fitzroy Ter., Prospect SA. 
t°33. Mitchell, Puof. M, L., ]£Sa University, Adelaide. 
193K. Moorhouse, F. W\, M.Sc, Chief Inspector of Fisheries, Flinders Street, Adelaide. 

1940. Mori-lock, J. A. T., 37 Currie Street, Adelaide. 

19.36. *Mountfor0. C. P., 25 First Avenue, St, Peters. Adelaide. 

1944. Mi:i(RW.t t J. W.. Kiiffineerina and Water Supplv Dent. Port Road, Thcharloii, S.A. 

1944. 2ftj#jfHft A. R, ( BA, 62 Shefiudd Street, Malvern, S-A. 

1945. *NokfHCon: ) K. FI„ R.Afir.Sc. A.I.A.S., 16 Carlton Street, Roscfield, S.A. 
1930- Ockenden. G. P., Primary School, WoodviUe,. S.A. 

1913; *Osl*ojw, Prof. T. G. B./D.Sc., Department of Botany, Oxford. England Council. 

1915-20, 1922-24; President, 1925 26; I'ur-Prcsident, 1924-25, 1926-27, 
1037. *Pakmm p L W.. B Sc, c/o Mines Diriment, Flinders Street, Adelaide, 
19*15. PArnsnv, G„ 68 Partridge Street, G!enelg, S.A. 

1929. 1'AiiLL, A. G. M.A., B.Sc., 10 Milton Avenue, Fullauon, S.A. 

192ft. >Prpr.R, C. S., i>.Se_. Waite Institute (Private Mail ftuj. Adelaide— Comuil, 1041-43; 

rice-President, 1943-45, 194o-47; Prrvdcut, 1945-4o. 
1047, Poykton, J. O., M.D., MA. Ch.R. M.R.CS., L.R.C.P., University of Adelaide. 

1925. *Pi<n<ooTT, Pkok, J. A., C.B.K., D.Sc, A.T.C., Waite Institute (Private Mail Rap), 

Adelaide— Vcrco Medal, 1938; Cmiir% 1927-30, 1035-30; l- r icc-Pr r xidenl t 1030-32; 
Preside?, 1932-33, 

1926. Price, A. G., CMG.. M.A.. Litt.D., F.R.G.S., 46 Pennincton 'Terrace, North Adelaide. 
1945. Pryor, L, D., M.Sc, Dip.Fnr, 32 La Pcrouse Street, Griffith, N.S.W. 

19.17. *Ratt, W. L., M.Sc, Medical School, University o\ Melbourne, Carlton N. 5, Victoria 

1944. Rickmay, D. 3.. B.A^r.Sc, C.S T/R, Division of Nutrition, Adelaide. 

1925. RrcHA»nsoN, A. E. V., C.M.G., M.A., D.Sc., 314 Albert Street, East Melbourne. 

W47- Rirjmi.. W'. R., B.Sc.. Si. Mark's College, Penninetori Terrace. .Vorth Adelaide. 

1047. Rtk. C. E., 42 WavmnuTh Avenue, (bandore, S.A. 

194'.. toomssn^, K. G.. B.Sc.. 42 Riverside Drive. Sudhurv, Ontario, Canada. 

1945. Rymill, J. R., Old Penola Estate, Fennla, SA, 

1944. *SAvr>Ans, Mis;; D. F., University of Queensland, Brisbane Queensland. 

1933. SrHNEumR, M, M.B., B.S., 175 >!orth "TtT„ Adelaide. 

1924. *Sf.cnii\ R. W, M.A., B.Sc., ErohvjeriniT and Water Sufply Department, Victoria 

S'fnare, Adelaide— Srcrrtnrv, 193CK35; Couiinl, 1937-3H; Vhc.Pwniait, I0.W-.30, 
1940-41 ; President, 1939-40. 

1946. ^Suinit. E. R., M.Sc. C.S.l.R. Diuision ot Jndiir4riul Chcimstrv, Box 4331, G.P.O. 

Melbourne, Victoria. 

1925. *Shea*.d, R, Port Elliot, SA. 

1936. *Shf-\w>, K. ( Fisheries Research Div-CSXK.. c/n Tn^tirute nf Agriruhnre. Univ., W.A. 

1945. Shp.phkt^, J. H.. B.Sr^.. RA., e/o Zinc Corporation, Broken IJill, X.S.W. 

1934. SiiTN-KFiELn R. C, SaUshnrv, S.A. 

19J2 Sr\T\mvns, H. VV\, 150 Fisher Sfrcct. Fullartoui, S.A 

193ft. *StMr5.o^. Mr?;. E. R-. M.Sc., Warland Road, Rurnside. 

1924. Stmp^ok, F. N. t Pirie Street, Adelaide. 

1<)44. Smitit, C. A. N.. B.Aer.Sr . Waite Institute .Privale Mail Ba^l Adelaide., S A. 

1941. Smith, T. Lavopord-, B.Sc., Department oi P ns t War Recon.vtrucliou, CinherT^ 

1941. Southi.:ott, R. V., M.B., B.S.. 12 Avenue Rnad. Unley Park, S.A. 

1930. SouTnwtX-u>, A. R.. M.D.. M.S. (Adel,), M.R.CP.. Wootoona Ter., Glen Osmond SA 

1947. .Spkcttt, R. I'. L , R.Sc., 15 Main Road, Richmond, S.A. 

I93fi. *Sparr;c. K. C. M-S^,. Mine? Denartment. Flinders Street, Adelaide, 

1947- Svohij-vi,,, M. EL, B.Sc., Denartmeut oi A^ticidture. Adelaide. 

1WS. *STEPnn\-?. C. G_, M Sc. Waite Institute (Private- Mail Bps-), Adelaide 

1935. Strtcicla^d, A G, M.AprSc, 11 Woolooiw Terrace, Glen Osmond, SA. C«uwri/, 

1932. Swaw. D. (.. "NfSc. Waite Institute HMVatt Wftfl Rag), Adelaide— Secretary** 
1940-12* I'i-^Prcuilcnf. 1946-47; President. 19/I7-, 


Date of 

1934. Symons, I. G., 35 Murray Street, Lower Miitcham, S.A. — Editor, 1947-. 

1929. *Tayu)r, J. K., B.A., M.Sc, Waite Institute (Private Mail Bag), Adelaide— Council, 


1938. *Thomas, Mrs. I. M., (nee P. M. Mawson), M.Sc, 12 Broadway, Glenelg. 

1940. Thomson, Capt. J. M., 135 Military Road, Semaphore South, S.A. 

1923. *Tindale, N. B., B.Sc, South Australian Museum, Adelaide — Secretary, 1935-36; 
Cmincil, 1947-. 

1945. TrvER, N. S., B.Agr.Sc, Waite Institute (Private Mail Bag), Adelaide. 

1937. *Trumble, Prof. H. C, D.Sc, M.Agr.Sc, Write Institute (Private Mail Bag), 

Adelaide— Cmincil 1942-1945; Vice-President, 1945-46; President, 1946-47. 
1894. ^Turner, A. J., M.D., F.R.E.S., Dauphin Terr., Brisbane, Qld. 
1925. Turner, D. C, Brookman Buildings, Grenfell Street, Adelaide. 
1912. *Wari>, L. K., I.S.O., B.A., B.E., D.Sc.. 22 Northumberland Avenue, Tusmore— Council. 

1924-27, 1933-35; Vice-President, 1927-23; President, 1928-30. 

1941. *Wark, D. C, M.Agr.Sc, Div, Plant Industry, CS.I.R., Canberra, A.C.T. 
1936. Waterhouse, Miss L. M., 35 King Street, Brighton, S.A. 

1942. Watson, R. H., Central Wool Committee Testing House, 572 Flinders Lane, Melb. ( CI. 

1939. *Weedinc, Rev. B. J., P.O. Box 51, Minlaton, S.A. 

1946. Whittle, A. W. G. B.Se.. Bridgewater, S.A. 

1931. Wilson, C. E. C, M.B., B.S., "Woodfield," Fisher Street, Fullarton, Adelaide. 

1946. Wilson, A. F., M.Sc., University of Adelaide. 

1944. Wilson, E. C, M.A., B.Sc. High School, Rcnmark, S.A. 

1938. *Wilson, J. 0. t CS.I.R. Division of Nutrition, Adelaide. 

1930. *Womersley, H., F.R.E.S., A.L.S. (Hon, causa), S.A. Museum, Adelaide— i/<?r<;r. 

Medal 1943; Secretary, 1936-37; Editor, 1937-43, 1945-47; President, 194344, Vice- 
President, 1944-45; Pep. Fauna and Flora Protection Committee, 1945. 
1944. *Womersley. H, B. S-. M.Sc. 43 Carlisle Road, Westbotirne Park, S.A. 

1944. Womersley, J. S., B.Sc, Lae, New Guinea. 

1923, *Wooi>, Prof. J. G. f D.Sc. Ph.D., University of Adelaide— Verco Medatf, 1944; 
Council, 1938-40; Vice-President, 1940-41, 1942-43; Rep. Fauna and Flora Board, 
1940- -.President, 194142; Council 1944-. 

1943. Woodlands, Harold, Box 989 H, G.P.O., Adelaide. 

1945. Wortitley, B. W.. B.A., M.Sc. A. Inst. P.. 6 Kensington Road. Wattle Park, S.A. 
1942. Zimmer, W. J., Dip.For., F.L.S. (Lon.), 22 Docker Street, Wangaraita, Vict 



[Generic and specific names in italics indicate that the forms described 

are new tn science.] 

aRnvena, xy Unit's, 

Abarys anuiitrodes 40 

A cacia en thy car pa t pingu i folia, com mil is, 

nudula. 20 
Acanthocephala, 13 
Additions to the Flora of South Australia, 

No. 44. J. M. Black, 20 
Adelaide Series, The,, as developed along" thti 

Western Margin of the Flinders .Ranges, 

IX Mawson 259 
Algal ecology, oi Marine Algae of Kangaroo 

Island. H. B. S. Womersley, 228 
Alloea xyl^chroa y 38 
-Jwphibiophilus eyerniae, 23 
AnufTsra xnlhochrna, 37 
Anckyhdicln Implodes, 3$ 
Anerastria tlwdorhros, 

clcpsiphronica r 30 
Atiipo pallescoti, 226 

Iscoschonaastia uromys, 10, pchymiptva-, 11 
Aneytosis fhiostieha 53 
Australian Acanthocephala No. 5. T. Harvey 

Johnston and S. J. Edmonds, 13 

Hakeriola t&sntaMPiuL<; t rnhia, 227 
Beckwith, A. C, t T. H. Johnston and: Larval 

Trematodes from Australian Freshwater 

Molluscs Part :XII, 324 
ftelicmclla gitexi, 218 
Black. J, M. Additions to the Flora of South 

Australia, No. 44, 20 
Burbidge, N. T. Key to the South Australian 

Species of Eucalyptus.. 137 

Catcremna cataxutfui, mcdioUnca % irptoptila, 
48, mclanomita, 49 

Cercaria ancyti, 324, 326; lopkosoma, 328, 329 

Charnockitjc and Associated Rocks of 
North-Western South Australia. A. F. 
Wilson. 195 

Crocker, R L, and J. G. Wood. Some His- 
torical influences on the Development of 
the South Australian Vegetation Communi- 
ties and their Bearing on Concepts and 
Classification in Ecology, 91 

Cychmcdusa daindi, 220 

Dickmsonia costvta, 221 

Early Cambrian Jellvfishes from the Flinders 

Ranees. R. C. S'pritfS, 212 
Jldutcaria flinrirrsi, 215 
Emmalocera crossospila. 37 
EphcsHa pelopis 40 
Eucalvptus— Key to the South A ustralian 

Species of: N. T. Burbidge, 137 
Eurvmelids from Australia and New Guinea. 

J. W. Evans, 225 
Eurymctoides soyerenxis, matuana, utfiro- 

bmtmea. 225 
EittrambiruJa yyTwwdactylct, 3 

Euageta arcstades, 47 

Eucopherodcs concineUa. homophaea, homo- 

ccxpna, schematica, 44, phaulopa, 45. 
Euzophcru alhicosta, 46, flattie osta-j ischnop®, 

arrhyihmopis t 47. 
Evans, J. W. Some New Eurymelids from 

Australia and New Guinea, 225 

Finlayson, H. H. On the Weights of some 

Australian Mammals, IS2 
F I indcrs Ra nges , The Adelaide Series as 

developed along the Western Margin of 

the: IX Mawson, 259 
Functional Synthesis in Pedogenesis. G G. 

Stephens, 1/jg 

Geology of the Jamestown District. South 

Australia. T. Langford-Smith, 281 
Granites and Granitisaliou. K. k. Miles, 54 
(Tri'7'ilk'O laubcllifrra, 2\ 

Jlomocosnma centres ticha, rhapto, 41 i'nry- 
leuca, contractu pclnstirta, achropasla, 
atechw, 42, lechnosenta, 43 

Hypoecbinorhynchus alaeopsis, 13 

Hypogryphia am&todcs, 53 

Tpoella davisi, 227 

Ipoidcs frnmoivtandata, mclakucac, 226 

Jamestown District, South Australia. Geology 

of the: T. Langford-Smilh, 281 
Johnston, T. H.» and Beckwith. A. C. Larval 

Trematodes from Australian Freshwater 

Molluscs Part XM 324 
Johnston, T. H., and Mawson, P, M. Some 

Nematodes from Australian Lizards, 22 

Kerlins. L. C. P. Pythium debaryanum and 
Related Species in South Australia, 252 

Key to i,he South Australian Species of Euca- 
lyptus. N. T. Burbidge, 137 

Lamprothammum, 318 

Lane Poole, C. EL, and Prescott, J A. The 
Climatology of the Introduction of Pines 
of the Mediterranean Environment to 
Australia. 67 

Langfnrd-Stmih, T. The Geology of the 
Jamestown District. South Australia, 2S1 

Larval Trematodes from Australian Fresh- 
water Molluscs Part XII, T\ H. Johnston 
and A. C. Bcckwith, 324 

Lioprosopa, dhvvchla, phauhdes, pclflpo, 
phafochifon 31, pachyr.a?u:h?a t sporadica, 
rhodinodrs, twiybeto, haploa, 32, thia- 
nwch hi, coJohcla. platyinoeh !a f iransccta, 
33, polios tie ha, rlutntisla, 34 


Lvchiiorhamnus. 318 

Madigan, C. T. Obituary Notice, 1 
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the Flinders Ranges, 259 
Mavvson, P. M.. and "Johnston, T, H. Some 

Nematodes from Australian Lizards, 22 
Miles. K. R. Pre-Cambrian Granites and 

Granitisatiou, with Special Reference to 

Western Australia and South Australia, 54 
Mu^tirave Ranges. Outrnockitic Rocks of ; 

A. i*\ Wi!son, 195 

Nematodes from Australian Lizards, T. IT. 

Johnston and P. M. Mawson, 22 
Xitellopsis, 314 

Oaitischtmgastia cava, 8 

On the Weights of some Australian Mam- 
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Ophel, J. L. Notes on the Genera Lychno- 
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Ophel, t [Wi and Womersley, H. B- S. Pro- 
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Qphida scarfs varmii, 23 

Papiliatwta fytfi, 223 

Pararharfinorhynchns ■mupilis, 15 

Pamlhrlav.ifros, ocdurac, 25 

Pedogenesis. Functional Synthesis in: C. C. 
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Phar> ncrndrn tiliquaVe 26. australc. 27 

Phycitidae, A Revision of the Australian, A. 
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Fhysaloptera antarctica. confusa. 24 

Pinus radiata 70, 80, 82; canadensis 72, 80, 
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Pneumnnema ti'iquae, 24 

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Prcseoll. J. A., and Lane Poole, C. F. The 
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Protochara, a New GeiJus of Characeae from 

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Protochara australis 311, iujlaia i'iA 
Pvthmm debaryanum and Related Speries in 

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Pythium debaryanum, ultimum, polymorph '>*«. 

vcxans 252-257 

RhadLnorhynchus pristis, 17 

Saluria stcrcochortla. petachvoxi, Zii 

Schnngastia fihUipi, 6 

Sprigg, R. C Karly Cambrian (?) Jelly- 
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Trissonca clytopa, 52 
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Turner, A. Jeffries. A Revision of tht Acfcr 

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Womerslev, H. B. S., and Ophel. 3. J,, Pro- 
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Wood. J. G-. and Crocker, R. h. Some Ffcis : 
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Johnston, T. EL,, and Edmonds, S. J.; Australian Acanthocephala No. 5 .. .. 13 

Black, J. M. : Additions to the Flora of South Australia, No. 44 20 

Johnston, T. H., and Mawson, P. M. : Some Nematodes from Australian Lizards . . 22 

Turner, A. J.: A Revision of the Australian Phycitidae, Part 1 28 

Miles, K. R, : Pre-Cambriau Granites and Grauitisution, with special reference to 

Western Australia and South Australia 54 

Prescott, J. A., and Lane-Poole, C. E. : The Climatology of the introduction of Pines 

of the Mediterranean Environment to Australia 67 

Crocker, R. L., and Woon, J. G. : Some Historical Influences on the Development of 
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Burbidge, Nancy T. : Key to the South Australian Species of Eucalyptus L'Herit .. 137 

Stephens. C. G. : Functional Synthesis in Pedoi 168 

Finlayson, H. H. : On the Weights of some Australian Mammals .. .. 1 S2 

Wilson^, Allan F. : The Charnockitic and Associated Rocks of North-Western South 

Australia. Part I, The Mnsgrave Ranges— an Introductory Account 195 

Sprigg, Rkg, C. : Early Cambrian (?) Jell from the Flinders Ranges. South 

Australia . . . . 212 

Fvaxs, J, W. : Some New Eurymelids from Australia and New Gain moptera, 

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Womerslev, H. B. S.: The Marine Algae of Kangaroo Island. I. A General Account 

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Keuling, L. C. P. : Pytluum debaryanum and Related Species in South Australia . - 253 

Mawson, D. : The Adelaide Series as Developed along the Western Margin of the 

Flinders Ranges . . . . . . . . . . . . . . . . . . 259 , 

Langfurd-Smith. T.: The Geo >town District, South Australia .. .. 281 

Sprigg, Reg. C. : Submarine Canyons of the New Guinea and South Australian Coasts .. 2% 

Womersley, H. B. S.: Tlie Marine Algae of Kangaroo Island. I. A General Account 

Western Australia 311 

Opuel, I. L. : Notes on the Genera Lychnothamnus and LamprothaLinium (Characeae) 318 

Johnston, T. Harvey, and Beckw ; Larval Tn 

Freshwater Molluscs, Part XI 1 ....